This weekend (May 2nd - 4th) sees the 2014 Lyme Regis Fossil Festival taking over a small portion of south Dorset and offering all manner of talks, activities, stands and experiences for palaeontological aficionados. My palaeoart exhibit, Palaeoartworks, is situated in the middle of the action at the Town Mill and will be open until the end of the festival. Sad as it is that the exhibit is coming to a close, I'm looking forward to its final days as - finally - I'm going to get to spend some time at the gallery itself, being on hand all day Saturday and Sunday to meet festival goers and - for the first time - selling small prints of my work. If all goes well, I may look into setting up a more permanent print service.

To whet your appetite, I've posted the 'rolling gallery' - effectively a slideshow of my recent palaeoartwork - above. This video has been running for a near-month in Lyme Regis alongside the framed prints to show artwork which there wasn't space and funding enough to show. Even if you've seen it, check it out again: this is a slightly updated version with some new images. One update includes a slide urging support for genuine palaeoartists over those who merely copy and derivative work from creative individuals. As discussed here, this practise is detrimental to the artistic and commercial success of palaeoart, and needs to stop. This won't be the last you'll hear of this issue around these parts.

Where there's a logo, there's a way.

But that's not all!

At 16:00 on Sunday I'll be leaving the gallery to give a talk on palaeoart in The Hub, situated on Church Street just around the corner from the Town Mill. The talk celebrates palaeoart as a wide industry rather than just prattling on about my own work, so we'll be covering what palaeoart is, how it began, and how to approach the palaeoart process. If all goes to plan, there should be time for questions at the end. Unlike the gallery, the talk isn't free, but it's as good as: tickets are £2.50 per person, or £2 concession (you can pre-book tickets here). Visitors to the Town Mill will get something of an insight into palaeoart methodology beforehand of course, thanks to the Palaeoart Case Studies left dotted around Palaeoartworks. Tasters of what we'll cover are shown below, including a short video showing the development of my recent Baryonyxpainting (eagle-eyed viewers may briefly spy a few Scott Hartman skeletal reconstructions floating about as reference material).

Last week I gave a talk on the significance and methodologies of palaeoart to attendees of the Lyme Regis Fossil Festival. The presentation was filmed with the intention of being posted online but, sadly, was not quite of sufficient quality to present elsewhere. With this in mind, I've had a go at re-recording my narration over the slides, which is now clearer (though not perfect - see below) and available to check out above. There's quite a lot of stuff covered in here: the early history of palaeoart; discussions of why palaeoart is important; soft-tissue and colour restoration, All Yesterdays, and a lot more. This means that the topics are, by necessity, covered fairly briefly, but further reading is suggested at the end if you'd like to know more. Be sure to whack the quality up to 720p to see the slides in their best quality.

If videos of this sort are of interest, there's potential for several more. Like a lot of folks who do outreach and conference circuits, I've accumulated a number of talks over the years which are delivered once or twice to limited audiences and then never see the light of day again. Popping them online seems like an obvious way to maximise their potential so, if you'd like to see more of this kind of thing, do let me know in the comments below. If they prove to be popular, I'll look into investing into more substantial recording equipment to improve audio quality.

That's all for now but, if the combination of videos and palaeoart tickles your fancy, be sure to check out the rolling gallery from my last post.

Inside every 29 year and 11 month year-old man is a 5 year old who only watches monster movies so he can produce little doodles of them afterwards.

After much hype, Godzilla (2014) has finally stomped into cinemas around the world. It seems to have been a divisive monster movie, with some taking its slow pacing and focus on human characters as a breath of fresh air in a stale genre, and others wishing it had the wham-bam-thank-you-mam approach of Pacific Rim. Personally, I'm in the former camp, and am especially glad the new Godzilla franchise has not started with two hours of giant monster wrestling. While Godzilla (2014) is not flawless, it takes a lot of risks and feels like a modernised 'classic' monster movie, and not the focus-grouped, merchandise-led fanboy service it could have been. I should add that I'm not especially keen on the last 59 years of Godzilla movies, much preferring the genuine drama and symbolism of the original 1954 movie to the decades of monster fights which followed. Because Godzilla (2014) is tonally closer to the original than the later escapades, I'm not surprised that people just wanting two hours of skyscraper-sized wrestling are feeling a bit disappointed. Anyway, there's a lot we could say about this, but that's not quite what I want to write about  here.

A big part of updating Godzilla has been the overhaul of Big G himself, as well as bringing new creatures to this franchise. These are exciting times for monster film fans because, for only the second time in its 60 year history, one of the most famous movie monsters of all is in for a complete redesign (we all know what happened the first time) and, of course, is now free of the constraint of 'suitmation' effects - the fancy term applied to the 'guy-in-latex-suit' approach Godzilla movies are (in?)famous for. A fun fact about the original 1954 Godzilla is that the now-inconic chap-in-suit technology was a compromise, the studio initially desiring stop-motion effects like those of American-made films King Kong or The Beast from 20,000 Fathoms. Toho, the studio which created Godzilla, had neither the money or expertise to execute stop motion effectively, so opted for using costumes instead. Despite using their second choice, the Japanese trailer for Godzilla triumphantly declares their special effects superior to anything the American movie machine was making at the time! And it certainly worked out for Toho: after a difficult teething process, the 'suitmation' approach worked well for the franchise and wasn't tinkered with outside of the much maligned 1998 Tristar Godzilla.

For 2014, Godzilla is a fully digital and, as we all know, relatively faithful to the original designs. It has, however, been altered in ways which would be difficult to execute if we were still watching a man in a suit. A lot of these changes, as well as the design of Godzilla's adversaries, were pretty neat because they tie into what we know about animal biology, scaling and functionality, and I get the impression that the guys behind this latest Godzilla - Legendary Pictures - put a lot of effort into making half-sensible creatures which biologists, biomechanicists and functional anatomists can be relatively happy with. And yes, yes yes: there's a buttload of stuff which is clearly nonsense: there's no way these animals could be the size they are, or firing beams of nuclear fire from their throats and so forth. But that's just par for the course for a Godzilla movie, and I'm not going to jump on boring old bandwagon of highlighting how impossible the whole lot is. What's far more interesting, and what I want to focus on here, is how Legendary built their animals around standard movie monster tenets to produce creatures which are not only intriguing and cool-looking, but also chime with real animal biology and functionality.

Twinkle toes

Godzilla's stubby, new-look foot, as seen from an airport terminal. Image from the Godzilla movie forum.

One of the most striking aspects of Godzilla’s redesign is its short, stubby feet instead of the long, plantigrade feet we’re familiar with from the suitmation costumes (above). This is easily one of the best changes to make in the entire redesign. Not only does it move away from Godzilla looking like he’s wearing a pair of comedy slippers, but it makes a lot of sense from a functional perspective. Long, plantigrade feet of bipedal animals (like our own, as well as those of chaps in Godzilla suits) lift our entire body weight with every step, our metatarsals and ankles both supporting and propelling us forward. This works fine for lightweight animals like ourselves, but comparable foot anatomy in a skyscraper-sized animal would need to be immense to avoid buckling under thousands of tonnes of bending force. The superficially sauropod-, tortoise- or elephant-like foot of the new Godzilla negates this risk however, creating a more columnar distal limb structure which is not employed in lift and propulsion but instead ideal for supporting terrific weight. This has several effects on limb structure and locomotion. The foot musculature (anchored to the shin) is reduced, keeping the weight of the limb down overall, thus making movement more efficient. However, it also decreases the foot mobility, limits stride length and overall gracility. Some of these effects are countered, however, by the elongate thigh region common to all Godzilla designs, which can swing the shortened distal-leg over great distances with every step. This increased length of the proximal limb region and shortening of distal is a trend we commonly associate with larger animals, even in species which are quite active and sprightly, and descended from cursorial ancestors (e.g. rhinos). Godzilla’s enormous thighs also fit with observations that proximal hindlimb musculature becomes relatively huge in larger bipeds - see the pelves of large bipedal dinosaurs for examples - providing the powerhouses necessary to move its huge bulk around. The result is a limb which isn’t going to win Godzilla any prizes for sprinting but, more importantly, is much more suited to the animal itself than the older designs. And let’s face it, when you can take strides measured in hundreds of metres, moving relatively quickly isn’t a problem.

Godgilla

Gills ahoy, on the lower portion of Godzilla's neck. Image from Detroit News.

A clear departure from all other Godzilla movies is that new version can breathe underwater. They’re easy to miss, but look closely and you can spot a series of fish-like gills on the side of its neck. At certain points in the movie, you can see them opening and closing as the big chap respires. Apparently, these were added not only because Big G spends a lot of its time swimming from location to location in the movie, but also to explain how an animal as large as Godzilla could keep a relatively low profile for so long. Air breathing animals of such size would surely be better known to the world at large if they were routinely surfacing to take breaths. These are all that's left of fish-inspired designs for the new Godzilla, which also featured fins instead of rows of bony plates; fish-like scales instead of reptilian skin and so forth.

Gills on a big, adult tetrapod sound a bit crazy, and I will admit - the fish-like (bony?) gills of the new Godzilla are difficult to rationalise entirely among real animals. If we’re willing to stretch belief a bit (I assume we are, what with a fictitious 100 m tall reptile being the subject of discussion here), we can find saving grace from early tetrapods and their descendants, modern amphibians. The latter have internal and external gills in at least their larval stages with most species losing them as they become adults. These gills are supported by a series of bones behind the skull which lead to openings in the body wall which allow swallowed water to escape - these are known known as gill clefts. Some amphibians - specifically select caudatans (salamanders, axylotyls, newts etc. - see below) - famously retain gill structures into adulthood, including their elaborate external gills. We know from exquisitely preserved fossils that such gills were present in the ancestors of all modern tetrapods, some of which also retained these gills in adulthood, and others which at least maintained their gill clefts (Schoch 2009). It seems likely, then, that the possession of gills, and possibly retention of gill structures into adult life* is the ancestral condition for all early tetrapods - including, presumably - Godzilla. Amniotes, which also presumably includes Godzilla, have obviously abandoned these structures and co-opted the gill apparatus for other purposes.

An axolotl with its external gills. © Kevin Schafer / www.photoshot.com, linked from ARKive.

Could Godzilla or other creatures ever get their gills back? Well, maybe. For a long time it was thought that evolution was irreversible ('Dollo's Law of Irreversibility') and, under this concept, loss of gills would be a one way street: Godzilla would have to evolve an entirely new set of gills, which is entirely possible. However, it's now appreciated that Dollo's Law is not really a law at all, and that organisms frequently do reverse their evolution, including backtracking on complex aspects of their anatomy and life history: digits, dentitions, wings, larval stages, reproductive strategies are all examples of things which have been lost and then recovered in animals. What once seemed like a law is likely more statistical improbability: a lineage can evolve in many ways at any point in time, and the likelihood that it will directly reverse along a familiar path is relatively low. However, in theory, there's no real reason why this shouldn't happen, so long as the selection pressures are correct. It would take some hefty tinkering with genes and mountains of heterochrony, and it's difficult to think of conditions which would promote the development of early-stage gill anatomy, but I guess it's not impossible for an amphibious creature to regain its ancestral gill condition under the right circumstances. My guess is that they would look more like the gills of caudatans than they would the scale-like fishy apparatus of Newzilla, but, let's face it, big feathery gills would look incredibly silly on a city-smashing monster.

See? Less "Godzilla, King of the Monsters", more "Preszilla, Queen of the Desert". Both are scary, but in very different ways.

Has Godzilla put on weight? No, it's just in it's head

One of the key comments made about the new Godzilla design is that it’s a bit of a heffer compared to other versions. This is debatable (they’ve always been pretty chunky if you ask me), but Legendary’s Godzilla design probably appears fatter than the rest because of its proportionally small head. There are good design reasons for this move: the smaller head gives a natural taper towards the top of the animal, distorting its perceived perspective and size, and it helps avoid the comically oversized, googly-eyed heads of previous versions. It's also in keeping with the natural world. Animals show a disproportionate reduction in skull length with respect to body mass - that is to say, larger animals generally have proportionally smaller heads than smaller ones. Note that this applies to carnivores as well as herbivores (Van Valkenburgh 1990; Christansen 1999). Godzilla represents a real extreme of animal gigantism, so it makes sense that its skull and head is going to carry this allometric trend to an unprecedented limit. What’s more, whatever Godzilla actually is (the new movie, thankfully, doesn’t really concern itself with this), it’s clearly on the reptile branch of the animal tree. Reptile cranial musculature is generally less developed than that of mammals, so their skulls and heads are relatively smaller at a given body mass than equivalently sized mammals (Christansen 1999). Again, this fits neatly with the small cranium of our new-look Godzilla. Of course, making the head look even smaller are the big legs and counterbalancing tail but, as mentioned above, this is also to be expected. Bottom line: far from being fat, 2014's Godzilla is just showing the extremes of proportion that we would expect if an animal ever grew to the ridiculous sizes we see on screen, and my feeling is that we're looking at a far more 'realistically' proportioned version that we've seen before.

"I'm not fat, I just have a head of predictably small proportion based on typical animal scaling allometries which distorts your overall perception of my size!" Whatever, buddy. Image from Godzilla Movies.

MUTO: Finally, a convincing giant, flying movie creature

Moving away from the main attraction: Godzilla vies for attention in the 2014 movie with another species of giant critter, termed ‘MUTO’ (below). The design for the MUTO(s) is very cool: gigantic, eight limbed, insect-ungulate-pterosaur beasts which defy easy classification. We’re shown pronounced dimorphism in adult MUTOs, the females being larger and more robust than the males, and devoting all limb sets to either terrestrial locomotion or gripping. The males, however, allocate one set of limbs to flight, bearing a spectacular pair of enormous membranous wings which recall monowing plane designs of the early 20th century (this isn't a coincidence: male MUTO wings are actually based on stealth bombers). These wings are largely unfoldable and situated on the dorsal surface of the torso so, when grounded, they extend behind the MUTO like a huge cape. Despite being the smallest creature in the film, male MUTO is still huge, so there’s little risk of it clipping its flowing wings on any structures when locomoting terrestrially.

MUTO in flight, from the Godzilla trailer.

There’s a lot to like about the male MUTO’s wing anatomy and flight mechanics. Firstly, the wings of this animal are simply enormous, as they should be for a creature of its size. Not only do they dwarf the body of the animal, but we could measure their span in metric villages. All too often, flying movie creatures are equipped with wings far too small for their body size, but - running off intuition here - these look ‘right’ for its size. As demonstrated by the birds below, wings increase in length and area disproportionately with body size because of standard rules of scaling: any linear size increases equates to a cubed increase in mass, which gives more for gravity to pull against and greater wing area requirements to achieve lift. If we are ever to expect a several-hundred tonne flying creature to become become and stay airborne, it would need significant wingage: MUTO delivers on this front.

A rule of thumb for designing flying animals: as mass increases, wing length increases faster. Demonstrated here by various bird planforms from Rayner (1988).

But it’s not just wing size that’s cool about the male MUTO: it’s flapping, or lack thereof, was also neat. Flapping amplitude - the degree of movement of a full wing stoke - decreases with wingspan in all flying creatures. The larger an animal is, the less movement is required in its wings to produce an effective flap cycle. A familiar example of this is the wing motions of flapping geese compared to pigeons: both flap their wings nearly constantly while flying, but the former move their wings around the horizontal, while pigeons swing them in great arcs. All too often, giant flying movie creatures (including several giant pterosaurs) show whopping huge flapping amplitudes well beyond necessity for flight and, probably, aerial stability. Godzilla’s male MUTO moves his wings just a little, however: he definitely flaps, but his wings move just a few degrees around the horizontal, not in huge arcs (watch the clip, below, for an example. Skip to 0:24 if you're a particularly impatient type). He also largely soars, as would be expected for any uber-large flier: flapping those wings would take a lot of effort, so soaring is the way to go. It would be loads of fun to get some basic wing size and mass data for this guy to work out some basic flight parameters - cruising speed, travelling distance etc. Given that giant pterosaurs seem capable of cruising at 100kph (Witton and Habib 2010), and yet are mere pipsqueeks at 10 m wingspans compared to the male MUTO, I reckon we’re looking at cruising speeds of several hundred kph. And that’s if he’s not in a hurry.


Finally, we get to see the MUTO launch a couple of times in the film, and each time we see something reminiscent of quad-launching. Pterosaur fans will know this trick as the best hypothesis on the table for explaining how flying reptiles became airborne, and a core factor explaining their attainment of gigantic proportions (Habib 2008). The kimeatics of MUTOs takeoff aren't quite the same - which is to be expected seeing as the limbs of pterosaurs are quite different to those of MUTOs - but it's neat to see the same basic components - a standing start, the rocking back crouch, forward thrust, upward push and leaping phase - incorporated into MUTOs takeoff. These actions are all performed with the four walking limbs rather than the wings themselves, as in pterosaurs and bats, but this also chimes well: MUTOs would need all the power they could to become airborne, so using every available limb is logical. Note that insects are an unlikely model for MUTO takeoff because, while seemingly comparable with MUTO because of their similar limb and wing configuration, at least some insects only use one pair of limbs (the mesothoracic) to launch (Trimarchi and Schneiderman 1995). This is probably because insects enjoy much better power/weight ratios than large vertebrates/skycraper-sized-monsters. 2-4 mm long fruit flies can jump up to 15x their own body lengths - about 30 mm - using their middle limbs only (Zumstein et al. 2004). This sets them apart from bats and pterosaurs, which use all their limbs in launching, and makes these the best candidates for MUTO launch models. It would be neat to know how much attention the animators and designers have been paying to bat and pterosaur research or if they came up with this launch strategy on their own. My experience of explaining quad-launch is that a lot of folks find it counter-intuitive at first, probably because we don't see many animals routinely taking off in this way, so I do wonder if the similarity between these takeoff strategies reflects research into animal takeoff.

And that's it for now. If you want to know more about how the animals of Godzilla (2014) were constructed, you should check out the concept art book which accompanies the film: Godzilla: the Art of Destruction (Cotta Vaz 2014). Closer to home, more fun with famous fictitious monsters can be found here, and for more on movie creatures - specifically those of special effects legend Ray Harryhausen, check out this.

References

• Christiansen, P. (1999). On the head size of sauropodomorph dinosaurs: implications for ecology and physiology. Historical Biology, 13(4), 269-297.
• Cotta Vaz, M. (2014). Godzilla: the Art of Destruction. Titan Books.
• Habib, M. B. (2008). Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, 159-166.
• Rayner, J. M. (1988). Form and function in avian flight. In Current ornithology (pp. 1-66). Springer US.
• Schoch, R. R. (2009). Evolution of life cycles in early amphibians. Annual Review of Earth and Planetary Sciences, 37, 135-162.
• Trimarchi, J. R., & Schneiderman, A. M. (1995). Initiation of flight in the unrestrained fly, Drosophila melanogaster. Journal of Zoology, 235(2), 211-222.
• Van Valkenburgh, B. (1990). Skeletal and dental predictors of body mass in carnivores. In: Damuth, J. and MacFadden, B. (eds). Body size in mammalian paleobiology: estimation and biological implications, 18, 1-205.
• Witton, M. P., & Habib, M. B. (2010). On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness. PLoS One, 5(11), e13982.
• Zumstein, N., Forman, O., Nongthomba, U., Sparrow, J. C., & Elliott, C. J. (2004). Distance and force production during jumping in wild-type and mutant Drosophila melanogaster. Journal of experimental biology, 207(20), 3515-3522.

From Titan Books.

The Paleoart of Julius Csotonyi: Dinosaurs, Sabretooths and Beyond (Csotonyi and White 2014) is another palaeoart-focused book from Titan Books, who brought us the acclaimed Dinosaur Art: the World’s Greatest Paleoart two years ago (White 2012). Anyone familiar with this book will immediately note the very similar format and high production quality in this recent Titan release. Although slightly smaller than Dinosaur Art, this will - again - leave readers wondering how the extremely affordable price (£25) covers production costs. Csotonyi was, of course, one of the artists featured in Dinosaur Art but, this time, is entirely running the show. Fans of his work will have little doubt that he can carry an entire book by himself. For the last decade Csotonyi has been establishing himself as one of the world’s premier palaeoartists, illustrating countless press releases, books, articles and museum walls with intricate paintings or digitally-manipulated photograph composites. His work is in such demand that he is one of the few individuals globally who can make a living out of palaeoart, a status which is testament to the quality of his work.

Before we get into the review itself, I want to stress how much of a milestone this book is. Palaeoart and palaeoartists suffer a PR problem where artists are considered unimportant and interchangeable: individuals who are secondary to the scientists pushing palaeontology forward and the audience who – often superficially – experience their work. Titan Books showed that palaeoart could be tackled more seriously and respectfully with Dinosaur Art and are cementing this idea in dedicating a whole book to a leading palaeoartists. Csotonyi's position as a working palaeoartist with major publisher support is rather exclusive, but exactly the sort of treatment palaeoart needs. I hope that Csotonyi’s solo album sells well enough to kickstart a series of books featuring other artists. Intentionally or not, Julius’ artwork is a good place to start this hypothetical series: aesthetically pleasing, extremely high quality, and blending traditional palaeoart approaches with some more complex and radical compositions. As a means to test the market for these sort of books, Julius is one of the strongest candidates currently available.

As an industy, palaeoart needs all the help it can get, starting with this logo.

Right, big-picture stuff out of the way now: what of the book itself? At its most basic level, The Paleoart of Julius Csotonyi is effectively an expanded version of his chapter from Dinosaur Art, juxtaposing imagery alongside an interview about Csotonyi’s art, influences and background. The interview, confined to the first 23 of the 156 pages, features intelligent questions and the interesting responses from the artist. Csotonyi’s passion for art and science are clear even before his images are displayed in earnest, as is the amount of work required to produce the large, ultra-high-quality imagery he is famous for. He leaves no doubt that many personal sacrifices are required to work as one of the world’s leading palaeoartists. This section also contains rarely-seen early works and non-palaeontological artwork, including some dedicated to astronomy. Some of the interview responses and other text features words which may be unfamiliar to lay audiences, but a glossary is provided to help readers navigate these terms.

The real meat of the book is relatively text-light so as to provide maximum space for Csotonyi’s art – large format is the only way to appreciate the detail it contains. The art is roughly arranged in chronostratigraphic order, with Palaeoazoic, Mesozoic and Cainozoic subjects separated into different chapters. As usual within palaeoart, the bulk of the artworks depict Mesozoic dinosaurs, and theropods are particularly well represented. Each piece is accompanied by brief details of the composition and commissioners, and some featuring additional comments from scientists about the subject animals. These comments mostly complement Csotonyi’s talents or spin yarns about research associated with the depicted species and, I guess, are designed to boost the scientific content of the book. I do feel a trick has been missed here because none pass particular comment on the decisions made when reconstructing the animals. Seeing as a lot of Csotonyi's art is produced alongside consulting scientists, I’d like to know what input they had. Even the most tightly constrained reconstructions of a fossil animal requires a lot educated guesswork and speculation about palaeobiology and life appearance and - in my own experience at least - not all of this is left to the artist. After all, this is a primarily a book about scientific art, and it seems that these comments could be more insightful than discussions about fossil localities, chance discoveries, or another complement for Julius' artwork (meant with all due respect, of course, but we know he's good. That's why we bought the book!).

A busy day in Permian Texas. Photo composite by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.

Csotonyi himself gives some works longer explanations about the processes involved in the reconstruction. These often highlight works with unusual compositions or viewpoints (such as the excellent ‘fish-eye’ sauropod view) and describes the way each piece was executed, often with alongside draft versions. These provide some insights into his process and will doubtlessly be useful to budding artists. My personal take-home message from these is the exhaustive consideration and research required to understand not only fossil animals, but to also reproduce realistic landscapes and lighting, particularly when odd perspectives and water are involved.

On the art itself: Csotonyi’s images are created using a range of media, including traditional and digital painting, sketches and – most commonly – digital photographic manipulation. I’m going to come clean here and admit that I’m not enormously fond of photographic manipulation. Many such works often fall into palaeoart’s own variant of the ‘uncanny valley’ or, all too often, present oddly-proportioned, strangely posed creatures which have little in common with their known anatomy. Julius’ photo composites are easily among the best, if not the best, attempts at photo-realistic 2D palaeoart out there however, and present reasonably reconstructed animals at either photo-realistic quality, or within inches of it. Some images, particularly the more ambitious, crowded scenes (fans of ‘a busy day in deep time’-type images are well served here) do bear niggles which jar the illusion, such as animals appearing too sharply defined against the background. To a certain extent, this is unavoidable: photomanipulation is incredibly difficult to pull off even remotely well, and even Csotonyi’s lesser successes are still amazing efforts. There are no overused photographic elements, no blurred skin textures, no cloning of animals to create herds of the same individual. When the photomanipulation does work well – and it frequently does – the effects are nothing short of stunning (e.g. below). The image of the resting Edaphosaurus on page 33 could easily be mistaken for a genuine, beautifully shot photograph. As with Dinosaur Art, some panoramic scenes unfold to show enormous vistas stuffed with detail. Many of these fold-outs allow those of us with empty pockets our first detailed look at the many murals Julius has created for North American and Australian museums.

Photo composite Acrotholus audeti and Neurankylus lithographicus  by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.

My favourite images in the book are digital paintings (below), such as the dancing Guanlong, the mothering polycotylid and the ceratopsid portrait gallery on pages 102-103. Not only do these show the trademark Csotonyi attention to detail but they’re wonderfully lit and composed: they feel more ‘of a scene’ than the photo composites. A neat touch is that alternative versions of well-known paintings are sometimes included. I actually prefer the near greyscale version of the Acheroraptor press release image on page 43 to the original, its dusky palate and the removal of the mammal from the hero animal’s mouth creating an entirely different tone to the more familiar version.

Digitally painted Brachiosaurus by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.

Of course, scientific accuracy is also essential for palaeoartworks to be considered successful. In this respect, the book also delivers. Thoroughly modern reconstructions of fossil subjects are presented: extensively feathered maniraptorans, diverse integuments in other dinosaurs, correctly orientated limbs and so on. In light of All Yesterdays (Conway et al. 2013), Csotonyi’s approach to extinct animal reconstruction may be considered conservative - there are no outlandish, speculative audacities here in terms of appearance or behaviour. Most of the depicted animal interactions are predatory, and the soft-tissues of the reconstructed species are not especially elaborate. Perhaps this is because nearly all of the artworks were commissioned by researchers and museums, clients who tend to favour safer, more conservative palaeoartworks. We should not lose sight of how progressive even ‘conservative’ modern palaeoart actually is. Many sights now familiar to us would have been considered heretical just a few years ago: Csotonyi shows several tyrannosaurs with variable amounts of feathering in the book with little fanfare, for instance. For dinosaurs at least, it’s becoming harder to produce wholly shocking palaeoart without unreasonably bending palaeontological science or speculating wildly. While Csotonyi’s book may lack the accessory frills, wattles and elaborate behaviours of some modern palaeoart, it acts as a fantastic milestone for how far palaeontology and palaeoart has moved in recent years. Moreover, I do not want to give the impression that the images are not interesting or novel: fishing Dimetrodon (above), Polycotylus nuzzling its offspring to the water surface to breathe and reptiles swimming between the dredging fronds of rafting crinoids are just some thought-provoking Csotonyian innovations.

The unkillable skim-feeding hypothesis lives on. Art by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.

I do have a few issues with some science behind the artwork. I’m told that an unfortunate misunderstanding resulted in the extensive discussion of Rhamphorhynchus skim-feeding on pages 136-139 (Hone, pers. comm. 2014; above. See the comment from Julius below for the sull story). This was, in fact, meant to reflect dip-feeding or surface-gleaning. Folks who keep up with pterosaur research will know that skim-feeding habits in flying reptiles has been looked into several times, consistently found problematic (e.g. Chatterjee and Templin 2004; Humphries et al. 2007; Witton and Naish 2008, 2013), and widely publicised. It’s a surprise and a shame, then, that this idea made it into the book without someone noticing, and particularly so because the science elsewhere is pretty tight. I also wonder if some of the photo composite crocodyliforms are shown with entirely accurate scute patterns, as most seem to have been taken from modern crocodylians – many Mesozoic crocs had very different, often simpler scute morphologies. And while we’re moaning, I do wonder if some more complex images would have benefited from small ‘key’ illustrations demonstrating the position of each animal. This is not only because the animals can be hard to spot in the complex, detailed scenes on offer (this is not meant as a slight – remember that many of the more complex images are intended to be hundreds of times larger on museum walls), but because linking a list of unfamiliar names to specific creatures can be difficult. Individuals intimately familiar with genera of all major vertebrate groups should be OK (they exist, honest), but I suspect they will only make up a fraction of this books audience.

These are only minor issues in the grand scheme of things, however. The intelligence and quality of The Paleoart of Julius Csotonyi makes it essential for anyone interested in palaeoart, as well as more general aficionados of palaeontology, natural history, or natural history art. I have no doubt that palaeoartists will be keeping a close eye on its success, and hoping that it presents the first of a wave of similar tomes from Titan Books. That’s all to come, though: for the time being, The Paleoart of Julius Csotonyi cements Csotonyi’s status as a world leader among the current crop of palaeoartists, and this book will only further his success.

References

Darwinopterus robustodens and his pal, a European robin (Erithacus rubecula). When not posing for artwork, they drive around in a van solving mysteries.

Wukongopterid pterosaurs have been on my mind this week thanks to a near-complete manuscript about them. This required* rendering everyone's favourite wukongopterid Darwinopterus - results above (note that this is D. robustodens, not the more familiar modularis). In the final version he's joined by something else, but you'll have to wait to just what that is. Wukongopteridae is the group of recently discovered Chinese pterosaurs which bridge early pterosaur and pterodactyloid-grades of pterosaur morphology, famously combining head and neck characteristics of the latter with the bodies of the former. They're best known for Darwinopterus modularis, but there's actually now nine(!) taxa known from the same horizon in north-east China, all morphologically very similar and almost certainly oversplit.

*By 'required', what I really mean is that I'm compelled to construct papers with extraneous artwork in them somewhere, because I'm a sucker for punishment.

Illustrations of Darwinopterus and I go way back. I was asked to do press images for two early publications on it: the initial description and assessment of its unusually 'modular' evolution (Lü et al., 2010), and the equally fantastic discovery of its sexual dimorphism and an egg-mother association (Lü et al., 2011a). Below is the first of these images, published in 2009.

Ah, 2009. Shrink wrapping, relatively light integuments and very low soft-tissue crests were still in fashion. Fun fact: Darwinopterus was referred to as 'Frank' before it was given a binomial, a nod to it being a Frankenstein's Creature-like mash of body parts.

Looking back on Darwinopterus 2009, I'm not enormously happy with it. This in itself isn't that unusual. Artists often look back with dissatisfaction with older works, but this has an additional reason for dissatisfaction: I never really agreed with the notion of Darwinopterus as an aerial-hawker of flying tetrapods, and I think this comes across in its execution. I outlined my basic concerns with this idea in Pterosaurs (Witton 2013):

Given that wukongopterids have provided an insight into macroevolutionary processes, filled a gap in pterosaur phylogeny, and present a very unique pterosaur bauplan, expectations may be high that their proposed foraging strategies will also be rather amazing. Fittingly, some have proposed that wukongopterids were pterosaur top guns, their newly evolved long necks and oversize heads being used to prey upon dinosaurs, other pterosaurs, and even gliding mammals in midair (Lü et al. 2010). Such acts would be rather remarkable because, with even a generous mass estimate, the biggest Tiaojishan wukongopterids would not weigh much over 300 g (extrapolating data from Witton 2008), which is about the same as a modern feral pigeon.

At that size, tackling squirrel- sized mammals or crow-sized dinosaurs on the wing would be a feat earning praise from even the hardiest modern raptors, and wukongopterid skeletons would have to be brimming with offensive weaponry for this purpose. Vertebrate-hawking birds are renowned for their talons, incredibly strong feet, robust skulls, and powerful beaks (e.g., Hertel 1995; Fowler et al. 2009), while bats that subdue large vertebrates in fl“ight are also armed with formidable teeth and powerful jaws (Ibáñez et al. 2001). These adaptations provide the means to immobilize their prey quickly and efficiently, and are obvious advantages for animals grappling with large prey while in “flight. Vertebrate hawkers are also powerful fliers that can chase down their quarry and, once immobilized, carry the prey to a safe spot to eat. Pterosaurian equivalents would therefore require equally powerful “flight musculature to permit the same tasks. Unfortunately for the Darwinopterus raptor hypothesis, wukongopterids do not possess any of these requirements. None of their appendages bear the chunky digits and talons ideal for subduing large aerial prey items, and their long, comparatively delicate skulls and unimpressive teeth are ill suited to this task. Nor, for that matter, do they have the expanded shoulder regions indicative of the powerful “flight muscles needed to chase and eventually carry their prey. With this in mind, raptorial pursuits look doubtful for wukongopterids.

Witton (2013), p. 141-142.

There's a lot more that could have been said about this, but you get the idea: Darwinopterus and chums were small (jackdaw-sized - see image at the top of the post), delicately built animals for which aerial predation seems counter-intuitive and unlikely. I'm not the only person saying this, either: Lü et al. (2011b) and Sullivan et al. (2014) also raise points against the aerial hawking idea. We could go so far as to to label wukongopterid aerial hawking as another example of an 'extreme' palaeoecology based on cherry picked characteristics rather than considering a full suite of functional data.

So, on reflection, we probably started on the wrong foot but, hey, what can you do? Working as an artist is quite different to being involved in research: ultimately, you're a guy with a paintbrush being told to illustrate someone else's idea, even if you don't necessarily agree with it (also see Csotonyi and White 2014). With the notion of aerial predation being raised in the paper, specifically as a possible explanation for the development of pterodactyloid head and neck features Lü et al. (2010), it was an obvious choice for the image. But we didn't help matters by walking into some of the cheesiest, silliest things stereotypes of palaeoart, primarily because we were trying to make a small, fairly inoffensive animal look like a skydiving, dinosaur-eating-badass. We tried several different takes on this, varying aerial and terrestrial prey, and compositions which upped the voracity, such as this:

You'll need to make your own whooshing jet fighter noises and 'pew pew' laser sounds.

Eventually death-from-above was decided over death from sideways. But a key issue to tackle was that that the proposed prey for Darwinopterus was the same size as the predator itself (Sullivan et al. 2014 go into this more), and it's obvious that Darwinopterus is not a fighting, wrestling creature. To make the image work at all we had to play liberally with animal sizes: the maniraptoran (loosely based on Anchiornis in the final version) is tiny, and the Darwinopterus is a huge, ferocious juggernaut. I'm not the only artist who used this trick: the disappointing pterosaur documentary Flying Monsters 3D also had their Darwinopterus plundering undersized theropods. After we tinkered with reality, we then started piling on the cheese: the maniraptoran was made more reptilian-looking, it's head twisted to stare into the Maw of Destiny and, of course, it's mouth open to 'NOOOOOO!!!' it's impending fate. The result is 'Meat Loaf palaeoart', the sort which resembles the worst kind of rock album covers more than nature.

Think I'm being daft comparing palaeoart to cheesy hard rock album covers? How many depictions of extinct animals have the exact sameposeas the titular bat here? See links below for more examples. From Wikipedia.

The result isn't awful, but I think I've had more successful collaborations with the same authors, just because this doesn't feel realistic at all. From the science to the composition, the whole thing just seems 'forced'. Don't get me wrong: I'm sure Darwinopterus could be terrifying instruments of mortality to the right prey (probably terrestrial invertebrates according to Lü et al. 2011b and Witton 2013), and I'm not belittling the drama or viciousness which can occur in the lives of small animals. But life - thank God - doesn't look like the cover of a Meat Loaf album. There are some fossil animals that can - just about - pull off Meat Loaf palaeoart, but it's just not possible to turn small, fluffy, unthreatening animals into monsters. When we do, they're more liable to look goofy than impressive. Despite this, we do it allthetime. I understand why we do - it's exciting and marketable, and feeds into expectation that Deep Time as a monster-filled fantasy realm - but it also reinforces the perception that palaeoart is unsophisticated and aimed at children. But there may be another way.

Palaeontologists: next time you have a new, diminutive animal you want illustrated, forget the monster: play the cute card. The chubby little Darwinopterus at the top of this post intuitively seems closer to the reality of this animal than Darwinopterus 2009: AwesomeoSuperKiller. It's proportions, fluffiness and posture are all accurate, and it only looks cute because, well, it probably was: it's a small fuzzball with an oversize head and mischievous grin. If the Internet's obsession with cats and baby sloths has taught us anything, it's that cute sells better than violence. Indeed, this image is one of the most popular things I've ever put on Facebook, and (at time of writing) it's was posted less than a day ago. The great thing about the cute card is that everyone wins: the artwork should promote research just as well or better than it's Meat Loaf variant, because it appeals to wider demographics. The artist gets to render an animal in a more realistic light without jumping through hoops to monsterise it, and the world gets a new picture to 'coo' at. A few folks might even start to appreciate extinct animals in ways they never could when they're always shown screaming and fighting.

And if nothing else, it will mean we'll never have to discuss Meat Loaf album covers here again.

May God have mercy on us all. (Wikipedia)

Mountains of respect to whoever can name the sources and years of each of these silhouettes. If your surname is 'Naish', you're not allowed to play. Title slide from my TetZooCon talk.

What are you doing on the 12th of July this year? If you're in London and you fancy a day of tetrapod-themed fun - and who doesn't? - you could do a lot worse than attending TetZooCon, a whole day inspired by the famous Tetrapod Zoology blog and its podcast. TetZoo really needs no introduction to anyone reading this, being renowned for mixing semi-technical zoological discussions of Recent and fossil tetrapods with speculative biology, fictitious creatures, cryptozoolgy and, well, whatever else takes the fancy of hosts Dazza Nash and Johnny Conners.

The full TetZooCon timetable of events is appropriately broad and covers dinosaurs, cryptozoology, conservation, speculative biology, primatology, wildlife watching and two regular stalwarts topics of this blog - pterosaurs and palaeoart. I'm taking reigns for the pterosaur aspect of the morning (you can get a sense of what I'll be talking about above - more details on this soon) as well as taking part in an interactive palaeoart event with two real artists - Bob Nicholls and John Conway. This promises to be great fun, and allows audience members an unusually good insight into palaeoart processes. I'll be selling prints of my work alongside other artists and merchandisers - the Palaeoplushies will be in town! - so bring your pennies for products you won't find anywhere else. And if that doesn't already convince you that there's something for everyone, the venue is none other than the London Wetland Centre, and delegates are free to wander around it all day.

In all, it sounds like it's going to be a terrific day, and I'm pretty stoked to be invited along. If this post tickles your fancy, and especially you'd like events like this to become regular fixtures in your calendar, support the event by spreading the word and grabbing some tickets. Looking forward to seeing you there!

Azhdarchids: also available in flying. Depicted animal here is based on Quetzalcoatlus sp., but no taxon in particular.

In just a few weeks the world will stop for TetZooCon, a one day convention of all things we associate with the famous Tetrapod Zoology blog and podcast - tetrapods real and scientifically-speculative from wondrous, charismatic fossil reptiles to deceptively interesting small, brown herpetofauna. If you’re reading this, you’re slap bang in the middle of the TetZoo demographic and I guarantee* you’ll have a good time. Tickets are available until Friday 4th July, and it’s all going down one week later - Saturday 12th. Get your place while you can, or forever live with the shame.

*Guarantee not guaranteed.

In a surprise move, my contribution to TetZooCon features pterosaurs. Specifically, I’m looking at the way one group of pterosaurs has made major ripples in the palaeontology pond in recent years - and not just scientifically. The changing face of pterosaur science is certainly interesting, but an equally intriguing, rarely told story exists on the popular face of flying reptile research. To whet your appetite, here’s an 'extended abstract' of my TetZooCon talk, giving some insight into what we'll be talking about in a couple of weeks.

PteroPop

Pterosaurs are not unfamiliar characters in popular culture. They have been mainstays of science fiction literature since at least 1874 (Jules Verne’s Journey to the Centre of the Earth), made the jump to the silver screen in 1925 (The Lost World), and since starred in uncountable stories of time-travel, lost worlds and Jurassic Park-inspired de-extinction fiction. Neither are they strangers to public education, from being part of Benjamin Waterhouse Hawkins’ 1854 Crystal Palace menagerie to modern appearances in £multi-million documentaries. The hundreds of years of popular pterosaurs have rarely showed much adherence to flying reptile science: their appearances, behaviours and lifestyles mostly reflect a shorebird-like 'pterosaur archetype' rather than the specific anatomy and habits of a once-living taxon.

One of the many curious things about early azhdarchid reconstructions is the head nubbin - a short, posteriorly-directed conical(?) crest on the back of the head. There is nothing like this known from any azhdarchid. Far from being an early mistake, reconstructions with head nubbins persist until at least 2000. Slide from my TetZooCon talk.

But all that is changing. One pterosaur lineage has overshadowed ‘generic’ popular pterosaurs to bring aspects of ‘real’ pterosaurology to the masses, and has even stolen some limelight from dinosaur celebrities in the process. Media as disparate as documentaries and comic books show these animals in (basically) anatomically correct forms, with accurate but atypical postures, and behaviours which are far removed from the idea of pterosaurs being ancient seabird analogues. The animals in question are, of course, the long-necked, toothless and gigantic flying reptiles, Azhdarchidae.

In some respects, the recent surge of azhdarchid pop-culture uptake is a bit strange. It is not, for instance, that azhdarchids are a newly discovered group. Far from it, their fossils were found by at least the latest 1930s or early 1940s; good remains were apparent the 1970s, and the concept of Azhdarchidae was formalised in the early 1980s. They’re not new to popular culture either, having hung around its periphery since the 1970s to be wheeled out as 'Largest Flying Animals Ever' on occasion. These early popular azhdarchids showed little uniformity in their reconstruction - maybe even less than other pterosaurs at that time. Most bore little resemblance to actual azhdarchid fossils, either anatomically of functionally (above): hugely elongate wings, longirostrine skulls with snub-noses, pin-heads, toothed jaws, short-necks, long necks with swan-like flexibility were all rendered in artwork from 1970 - 1990. Art produced in the 1980s - 2D work by Greg Paul, Paul MacCready’s 1985 glider (below), and a (largely sculpted) azhdarchid skeleton mounted by the Texas Memorial Museum - were probably the first works to strike close to reality, but they’re still a bit short of the mark. John Sibbick’s better known and more influential 1991 snub-nosed Quetzalcoatlus was a step back from these more accurate works, accidentally making a chimeric azhdarchid from at least two Javelina Formation azhdarchids (this ‘snub nose’ almost certainly belongs to an unnamed, short-skulled azhdarchid from the same horizon as Quetzalcoatlus).

The famous 1985 'QN' pterosaur, a half-size gliding Quetzalcoatlus northropi and friends, including its designer, the late Paul MacCready (right of middle, in the tie). The model flew successfully multiple times and isn't a bad rendition of an azhdarchid, although many assumptions made in its construction conflict with modern pterosaurology. From MacCready (1985).

Why all the confusion over azhdarchid appearance? Most azhdarchid material known until the late 1990s was either too scrappy to inform artists about life appearance, while the more complete material (the small Quetzalcoatlus species) was infamously not published (still isn't!). The world at large was therefore not able to appreciate azhdarchid anatomy, so any artwork of them required more guesswork than usual. In some cases, entire 'reconstructions' were products of imagination. Not aiding the murky early phase of azhdarchid palaeoart was the transforming nature of pterosaur science which, in the 1980s and 90s, saw much of what we thought we knew about these animals turned on its head. Thus, artists who wanted answers to simple questions like standing postures, wing membrane attachment and so forth weren't always presented with straight answers. 1997 saw a potential change for the better when Unwin and Lü (1997) reclassified the Chinese Maastrichtian ‘nyctosaurid’ Zhejiangopterus linhaiensis as an azhdarchid, but few paid attention to this obscure species when reconstructing 90s azhdarchids, and artwork continued to remain of variable accuracy. The azhdarchid fossil record has not improved fantastically since 1997, only expanding via isolated, scrappy bits and pieces. Their sudden popularity and uniformity of reconstruction has nothing to do with a significantly improved azhdarchid fossil record, then.

Azhdarchids: over 40 Megafonzies of cool!

So, if azhdarchids aren’t new, and they’ve not sent a burst of insightful fossil material our way, why are they now so popular? Perhaps recent reappraisals of their appearance and behaviour have more influence here than anything else. Reconsideration of azhdarchid mass estimates (e.g. Paul 2002; Witton 2008; Henderson 2010; Sato et al. 2010), re-interpretations of lifestyles (Hwang et al. 2002; Witton and Naish 2008, 2013; Carroll et al. 2013) and flight characteristics (Habib 2008, 2013; Witton and Habib 2010) have recast azhdarchids from billboards of flighted animal size to muscular, terrestrially-competent predators and powerful fliers which were also giant. This has seen azhdarchids landing ‘major roles’ in palaeo pop media. In the last five years, erect-limbed, terrestrially stalking and quad-launching azhdarchids featuring in the BBC documentary Planet Dinosaur, Atlantic Productions’ Flying Monsters 3D, the 20th Century Fox film Walking with Dinosaurs 3D, the 20,000AD comic series Flesh and recent comics of Teenage Mutant Ninja Turtles, been made into at leasttwo figurines by CollectA, in the upcoming, Steam-released multiplayer game The Stomping Land, in Nathan Carroll's wearable pedagogic puppet form and even a rap. This uptake of the same pterosaur lineage is all the more surprising when you consider the diversity of influences and goals of these projects, as well as the near-infinite sea of fossil species which could take their place. More remarkably, these depictions of azhdarchids aren’t anatomically bad or variable, either: they have large, pointy heads with posteriorly placed crests, long necks, short wing fingers and long limbs. It seems azhdarchids have genuinely penetrated the pop-palaeo zeitgeist.

Azhdarchids in recent comic books, including Teenage Mutant Ninja Turtles and the fabulous 20,000AD series Flesh. Note the far right panel - terrestrially stalked TO DEATH!!! Awesome stuff - should really cover Flesh in more detail here some time. Slide from my TetZooCon talk.

The times, they are becoming very different

The uptake of azhdarchids in pop-culture may reflect their recent recasting as stylish, dominating predators of small, defenceless animals, a formula known to strike a nerve with the public - ask H. G. Wells. Do our palaeo pop-culture icons attain iconic status because, like some literary ones, they simply evoke cool characteristics and styles?

Perhaps more importantly, azhdarchids have - for what seems like the first time - persuaded popular culture to widely depict actual pterosaurs rather than an anonymous set of wing membranes and toothy jaws. That's pretty neat, as it means we're starting to break the notion that different pterosaurs are just minor variants on the same basic animal. I wonder if this applies to pterosaur science too, as research into azhdarchid lifestyles and habits is providing compelling evidence of palaeoecological variation within the group: shorebird, fish-eating habits applied almost universally to Pterosauria just doesn't work for these guys. With it being increasingly obvious that azhdarchids were doing their own thing, it's easier to start seeing other pterosaurs as potentially having distinctive lifestyles as well. Azhdarchids may be the thin end of the wedge in this respect for both popular and scientific circles.

Of course, no-one can predict how long our current interest in azhdarchids will last, nor what will happen to hypotheses concerning terrestrial stalking, quad-launch and so on. My gut feeling is that these ideas will stand up to scrutiny, but we can never predict what the fossil record or new studies will tell us. Whatever happens, these ideas and the animals they concern have gone some way to superseding generic ‘pterodactyls’ in palaeontological culture, replacing them with a more accurate and detailed appreciation of pterosaur diversity. But what next for azhdarchids? What advances in azhdarchid science are on the horizon? How might these impact their portrayal in popular culture? For that, you’ll have to attend TetZooCon and my talk. Tickets!

References

Last Saturday hosted an event which might, in future years, be considered a strange experiment. Set up in typical convention manner with attendance fees, invited talks and interactive audience activities, its unique selling point was its inspiration: a (largely) technical science blog which covers obscure animals in as much, often more, detail than you'll find in any textbook or scientific paper, as well as arcane topics such as speculative biology, natural history art memes, and cryptozoology. I'm talking, of course, of TetZooCon 2014.

NB. Like a chump, I didn't take a single photograph the entire day, so you'll have to make do with a very bland blog post.

Held at the London Wetland Centre on the 12th of July, TetZooCon 2014 was the latest expansion of the 'TetZooVerse', an internet enterprise founded on three incarnations of the Tetrapod Zoology blog and, more recently, a podcast, two internetcomics and on-demand merchandise. The brainchild of Darren Naish and (more recently) John Conway, it's undoubtedly one of the longest running and most successful science outreach exercises on the internet, and notable for covering complex narratives and scientific problems in the world of tetrapod studies. TetZoo fans thus comprise not only casual internet surfers but also researchers and practising scientists. Few other blogs can boast such appeal and far reach, making TetZoo one of the few internet enterprises which might manage the tricky move from the 'free' virtual world to one of admission fees, travel expenses and conference overheads.

In many respects, TetZooCon almost felt like watching a live version of the blog as different talks - essentially 'live blog posts' - covered an array of TetZoo-relevant topics. Unlike TetZoo, the floor did not solely belong to Darren and John, but shared by a host of excellent speakers. I'm not going to cover the talks in detail here because others havealreadydone so, but the topics included speculative zoology, amphibian conservation, wildlife photography, vertebrate palaeontology, crytozoology and mythical animals. Regular readers will know that I was among the invited speakers and covered changing perceptions of azhdarchid pterosaurs. As with other elements of the TetZooverse, these talks meandered from pure science (sauropod neck length) to almost humanist topics (mermaids, the cryptozoological leanings of Shakespeare). Many struck ground between these extremes, noting the interplay between science and culture and how they've influenced each other - for better and worse. Arguably, providing a platform for such talks and the diversity of topics was TetZooCon's greatest success. I've not been to a conference where talk topics varied so considerably and, in contrast to conferences with homogenous themes, there was no chance for getting subject-weary here. The talks were presented at pitch-perfect semi-technical level, assuming that the audience was intelligent and would have some prior knowledge of the broader subjects at hand (e.g. there were no, or only very brief, explanations for what things like Orang Pendek, sauropods or azhdarchid pterosaurs are), while also appreciating the room was not full of experts. It helped, of course, that the speakers and presentations were excellent. I definitely walked away with a greater education than I walked in with.

Other events included a palaeoart workshop, where attendees - led by palaeoartists John Conway, Bob Nicholls and, er, me - attempted to restore the life appearance of the historic 'Mantell Piece'Mantellisaurusfossil, and a TetZoo-themed quiz. The former was of interest for not only palaeoart aficionados, but also anyone wanting to know how fossils are interpreted. There was discussion over bone identification, how many individuals were represented by the specimen, how we could deduce the affinities of the animal and so-on, and we all compared images and notes at the end. The work of the lead artists was beamed onto the screen behind us so audience members could not only see what we were sketching, but engage in discussion with us about specifics of the fossil. Suffice to say (cheating ne'er-do-wells aside who recognised the specimen and simply drew an ornithopod), there was virtually no reconstruction consensus. Tours of the wetland centre and the obligatory pub dinner followed, while merchandise - including prints, 'official' TetZoo products and the much lauded Palaeoplushies - was on sale all day.

Was the event a success? As a speaker and delegate, my opinion is an unreserved 'yes'. There were enough delegates to generate that 'real' conference feel, the day was varied and interesting, and it was a lot of fun to be part of. With strict scheduling, custom 'palaeoart cams', delegation packs and almost flawless audiovisual performance (except for my own talk!), the day was pulled off with the sort of professionalism you'd expect from a long running conference rather than a first-time event. Most importantly, the day felt fresh and different from other conferences. As millions of TetZoo readers and listeners attest, there is a large audience for the 'offshoots' of zoological science such as palaeoart, speculative zoology and so-on, but few venues exist to chat about these topics outside of the internet. TetZooCon is a welcome plug in that gap.

Of course, whether we'll see a second TetZooCon depends on the transformation of online (and free) participation in the TetZooverse to financial and time commitments from potential delegates. This point is really why I wanted to write this short post. Turnout for this first event was good and, happily, conference overheads were recouped. At the same time, I don't think Darren and John slept in beds of gold leaves that evening. Events like these live and die on the whims of potential delegates so, if you were 50:50 about attending this time around and decided against it, rest assured that it was a blast and you won't want to miss out again. If the event passed you by entirely, but you like the idea of an annual celebration of the palaeontological and zoological blogospheres, then you'll also want to get on board next time around. This is a conference with lots of potential and, with enough support, it could become one of the most accessible, unique and interesting fixtures of the conference calendar. If it happens, I'll be booking my place for TetZooCon 2015 as soon as I can next year. If that's not high recommendation, I don't know what is.

Admit it, whatever you drive to work seems a little less adequate now.

Pterosaurophile Mike Habib was recently featured in a Scientific American article about the utility of pterosaur research. Let's face it, as cool as pterosaurs are, it can be hard to justify research into them when the world is faced with real problems like climate change, overpopulation, an enormous biodiversity crisis and Michael Bay movies. But Mike's interest in pterosaurs principally concerns biomechanics, quantifying the mechanical properties of pterosaur anatomy and seeing what it was capable of, and this sometimes allows transference of their evolutionary solutions to our own technological problems. Among other things, pterosaur biomechanics might be applied to some big projects: developing unmanned vehicles - including some which may explore other planets - and developing wind-stable fabrics. The latter may not sound very exciting, but wind-resistant fabrics are essential in all sorts of extreme activities, from exploring remote corners of the world (think tents), lightweight aircraft (parachutes, hang gliders, etc.) and extreme sports (wingsuits).

But that's small fry compared to one idea mentioned in the article. As part of an international team - including myself - pterosaurs may be launching air travel in a whole new direction. The manner in which pterosaurs took off - so called quadrupedal launch - offers a solution to a problem faced thousands of times around the globe each day: launching aircraft into the air as effectively as possible. As we all know, three lines of evidence point to pterosaurs launching quadrupedally, with most effort coming from their forelimbs. 1) animals launch using from their 'default' gait, and pterosaurs were quadrupeds; 2) pterosaur forelimbs are much more developed than their hindlimbs, whereas the opposite is true in hindlimb launchers and, 3) above a certain size, pterosaur hindlimb bones would actually fail in launch (Habib 2008, 2013; Witton and Habib 2010). These point to a powerful, quadrupdal launch mechanic which permitted even the largest, 200-250kg pterosaurs to take to the skies from a standing start, while birds - with their hindlimb launches - are seemingly capped at 70-80kg.

It's not only large birds which look enviously on pterosaurs. Most of our own aircraft require runways for takeoff. Vehicles which can take off without runways, like helicopters, are constrained to large size because of their power requirements and required wingspans. All aircraft launches require lots of fuel, and lots of space. It's unsurprising, then, that quad-launching giant pterosaurs have attracted the attention of engineers, as they clearly evolved a method of launch which is not only space and fuel-efficient, but also incredibly powerful. Practical results are undoubtedly years away, but the notion of a small, solo-pilot aircraft being capable of quad-launch and powered flight is realistic enough that we're seeking money for a project to test the waters. The concept we have in mind resembles a suit more than a plane - as Mike put it on Twitter, "think Batman x Iron Man" - alluding to concepts of the craft being controlled by a person strapped within the chassis, sort of like wearing a multi-million dollar pterosaur costume.

A visual history of pterosaur-inspired flying machines. 1, Ernst Stromer, 1913, basic glider model of Rhamphorhynchus wing membranes; 2, Hankin and Watson, 1914, a model based on their pioneering studies of Pteranodon flight (Hankin and Watson 1914); 3, Erich von Holst, 1957, a rubber band powered, wing flapping Rhamphorhynchus glider; 4, Cherrie Bramwell and George Whitfield, 1974, 7m wingspan Pteranodon glider based on their seminal 1974 paper; 5, Bramwell and Whitfield, 1984, half scale 4.5m wingspan Pteranodon made for the BBC; 6, Paul MacReady, 1984-85, 5 m span Quetzalcoatlus remote controlled, computer balanced glider (see MacCready 1985); 7, Margot Gerritsen, 2005, scaled Anhanguera with fully articulated wings built for National Geographic; 8, Matt Wilkinson, Rodger Highfield, and Vivian Bock, 2007, wind tunnel model of Anhanguera used to test Wilkinson’s hypotheses on pteroid orientation, 9, PteroFlight, our new project looking into pterosaur wing performance and its applications. Image compiled by Iain McCreary, used with permission.

What might such a thing look like? Sadly, it's not going to look like the thing at the top of this post. What you've got there is food for thought rendered by someone who's aircraft design skills boils down to watching science fiction movies, and who's engineering protocols are determined by Cool Points. It takes the idea of a 'pterosaur exoskeleton' to an extreme definition, right down to the limb proportions, wing folding and ability to walk about on all fours. Undeniably cool looking, just not very practical. But technologies and ideas taken to an extreme in this painting actually do exist. Augmentation of human frames with robotic exoskeletons is an intensive area of research and already employed to aid physically disabled people, as well as boosting the carrying strength of ground troops. Computers capable of flying deliberately unstable and responsive aircraft -manned or unmanned - are widely utilised. Large, controllable pterosaur-inspired vehicles with moving, adaptable wings have been researched for 100 years (above) and achieved flight (albeit not launch) on numerous occasions, with recent models featuring automatic computer control. The basic elements of this project - essentially a computer-supported, pterosaur-inspired lightweight flying exoskeleton - are at the far end of known technological spectra, not fantasy and hokum.

Of course, we're not going to see pterosaur-inspired suits catapulting people skywards tomorrow. Some serious research and developmental work is required before we see anything like a working concept or even - if we're honest - if it's possible at all. At this stage, however, this ultimate application of pterosaur research is not being ruled out. In other words, keep watching the skies - and check out Mike's Scientific American feature for more details.

References

Thalassodromeus sethi after some worms. Note: not a turtle. From Witton (2013)

Today sees the publication of an article challenging an exciting claim made in recent pterosaurology (Grellet-Tinner and Codrea 2014). If you missed it, the article concerned identifies a thalassodromid pterosaur in uppermost Cretaceous rocks of Romania and the erects a new species, Thalassodromeus sebesensis Grellet-Tinner and Codrea, 2014. At the centre of this is ODA-28, an (alleged) fragmentary cranial crest only fully exposed on one surface. None of this may not sound like a big deal, except that other thalassodromids - including the alleged sister species, Thalassodromeus sethi - are only known from the Lower Cretaceous Araripe Group of Brazil. T. sebesensis thus is about 40 million years out of time and thousands of miles out of place, and also occurring when azhdarchid pterosaurs basically represent the entire diversity of Pterosauria (Grellet-Tinner and Codrea 2014). Suddenly, the routine act of naming of a new animal is rewriting our understanding of pterosaur evolution.

There's more. Despite having only a scraps of bone to work with, Grellet-Tinner and Codrea (2014) suggested the T. sebesensis crest anchored muscles to form a ‘sizeable fleshy crest’, acted as a rudder in flight, that it somehow highlighted co-evolution between Romanian pterosaurs and angiosperms, and ecological segregation between azhdarchids and thalassodromids. All of these ideas are pretty radical in one way or another, especially considering the fossil material they are based on.

Blah blah blah… extraordinary claims, extraordinary evidence etc. When T. sebesensis was published it raised the collective eyebrows of pterosaur workers for all the wrong reasons. ODA-28 has no obvious ties to Thalassodromidae (or Thalassodrominae, if that’s how you roll - see Witton 2009), Pterosauria, or even to a cranial crest. Today, I and 19(!) other authors have said this in print (Dyke et al. 2014), noting that ODA-28 lacks any pterosaurian synapomorphies or even features typical of the group. As anyone who has handled pterosaur fossils can attest, pterosaur remains are distinctive at gross and microscopic level, and ODA-28 lacks any features expected in pterosaur bone (e.g. extremely thin bone walls separated by trabeculae). Any resemblance to the Thalassodromeus sethi holotype is entirely superficial, and shared characters between the two specimens - notably the ‘fossae’ at the base of the ‘T. sebesensis’ crest - are really incomparable on detailed examination. A clear lack of symmetry in ODA-28 shows it is not a medial skeletal element either, and thus not the cranial crest of anything. In short, cancel the text-book revisions: the temporal and palaeobiogeographical anomaly of ‘Thalassodromeus sebesensis’ is just a fairly major misidentification of a scrappy fossil (Dyke et al. 2014).

The 'flying turtle': the holotype of 'T. sebesensis' compared with the plastron of the turtle Kallokibotion. A, NHMUK R4930, the lectotype plastron of Kallokibotion magnificum with the portion corresponding to ODA-28 outlined in black (photo supplied by S. Chapman, Natural History Museum, London); B) ODA-28 (modified from Grellet-Tinner and Codrea, 2014). Abbreviations: hypo, hypoplastron; hxc, hypoplastron-xiphiplastron suture; ihc, intra-hypoplastral suture; ib, inguinal buttress; ps, pubic scar; meso, mesoplastron; mhc, meso-hypoplastral contact; pll, posterolateral lip; xiphi, xiphiplastron. Scale bar for A equals 50 mm. From Dyke et al. (2014).

In all, a bit of an anticlimax. How did our short paper end up with 20 authors? The response was started by experts in the terrestrial faunas of upper Cretaceous Romania, who asked me and the Natural History Museum’s Lorna Steel if we could contribute a few paragraphs targeting the flawed pterosaur identity of the specimen. While we were working, it emerged that pterosaur experts from Brazil were also planning a response. The editors of Gondwanan Research, who published Grellet-Tinner and Codrea (2014), understandably only wanted one response, so the two teams joined forces. By the time experts in turtles, Romanian fossils and pterosaurs were all on board, we ended up with a truly international background: the USA, UK, Brazil, Romania and France are all represented.

A final note: this is not the first time thalassodromids have been pulled to the top of the Cretaceous. Kellner (2004) and Martill and Naish (2006) argued that a partial skull and mandible from the Maastrichtian Javelina Formation of Texas represented a thalassodromid based on perceived similarities with the thalassodromid Tupuxuara. While others have argued against this idea (the mandible and premaxillary morphology are more similar to those of azhdarchids - Lü et al. 2008; Witton 2013) - these claims have not been met with a sledgehammer response because the suggestions are not unreasonable. Sure, I don’t think the Javelina material in question is thalassodromid, but I can see why others might. ‘T. sebesensis’ has been swiftly rebutted by a crowd of experts because the underlying science is so clearly bogus that all concerned with pterosaur and Romanian palaeontology wanted it’s impact nipped in the bud. A response from Grellet-Tinner and Codrea will be published soon, so we'll see what they make of our rebuttal. To end on a high: there are exciting pterosaur remains coming out of Romania, and some of them are in the review/publication system already. Hopefully, we'll have some news on these out soon.

References

Yesterday, a huge team of authors called out the science behind 'Thalassodromeus sebesensis', an alleged new pterosaur species 40 million years and thousands of miles out of time and space (Grellet-Tinner and Codrea 2014). As with many outlandish palaeontological claims, the evidence behind 'T. sebesensis' really falls apart rapidly under scrutiny, principally because the alleged pterosaur remains actually represent an unremarkable piece of turtle plastron (Dyke et al. 2014).

A, the plastron of the fossil Romanian turtle Kallokibotion magnificum, compared with B, the alleged holotype 'cranial crest' of 'Thalassodromeus sebesensis'. For further details, see yesterday's post.

Since then, the response to our comment has been published (Codrea and Grellet-Tinner 2014). I'll admit to being surprised that Codrea and Grellet-Tinner maintain the specimen as a pterosaur, and consider the arguments raised against our points as weak, hypocritical and problematic, but whatever: the two arguments are now out, and the palaeontological community can judge for themselves. CT scanning is apparently planned for the specimen (Codrea and Grellet-Tinner 2014), which should put 'T. sebesensis' to bed once and for all.

This post isn't really about that, though: it's about correcting a mistruth in Codrea and Grellet-Tinner's response. Their comment shows little decorum or professionalism, attempting to undermine our response with ad hominem potshots at some authors of Dyke et al. (2014), including criticism of their editorial skills and the taxonomic confusion surrounding specimens described by the authors. Moreover, they criticise us for not examining the specimen, UBB ODA-28, before publishing our response. They state that:

"...UBB ODA-28 is housed in an official and recognized Romanian institution, thus available for examinations to anyone interested. This includes Dyke’s July 2nd 2014 written request to examine UBB ODA-28, which was immediately granted, although, Dyke went on writing its hasty comment without examining UBB ODA-28."

References

The Triassic ornithodiran Scleromochlus taylori depicted as a nocturnal desert-specialist with filamentous insulation, fuzzy feet for purchase on drifting sands and a saltatorial means of locomotion.

Like actors with one famous character, fossil taxa can become typecast to specific ‘roles’ in palaeontological discussions. One fact of their palaeobiological significance is entrenched so deeply that they are seldom mentioned outside of this context. Examples include Archaeopteryx as the first bird, Mei as the cute sleeping dinosaur, and Darwinopterus as the bridge between major stages of pterosaur evolution. Packaging these animals into simple factoids obscures much of their other interesting palaeobiology, so we rarely hear about their other remarkable features.

Step forth Scleromochlus taylori, a small Triassic archosaur from the Upper Triassic (Late Carnian) of Scotland. For 100 years Scleromochlus has been implicated as a relative of pterosaurs (e.g. Huene 1914; Padian 1984; Gauthier 1986; Sereno 1991; Bennett 1996; Hone and Benton 2008; Brusatte et al. 2010, Nesbitt 2010) or, at very least, an ornithodiran representing a very early stage of stem-bird evolution (Benton 1999; Hone and Benton 2008).* This is about all we ever hear about Scleromochlus, however: nothing more than a milestone in the evolution of pterosaurs or dinosaurs. I'm guilty of it too: in my own book, Pterosaurs (Witton 2013), Scleromochlus just formed an anchor for discussing ornithodiran evolution. Undoubtedly, this needs correcting: Scleromochlus is a unique and interesting animal in its own right, and one fully worthy of detailed discussion. To relieve my shame, I'm going to attempt such a discussion here. Just for fun, I'm going to write it in the same style as a Pterosaurs chapter.

*You can't mention Scleromochlus on the internet without someone pointing out that its status as an ornithodrian has not been tested in analyses containing non-archosaur archosauromorphs. This is true enough, but - at least within the current limits of testing - its ornithodiran status is not controversial, having been recovered in at least six different analyses (e.g. Gauthier 1986; Sereno 1991; Bennett 1996; Hone and Benton 2008; Brusatte et al. 2010) and sharing several unique characteristics with Pterosauria (Padian 1984). Hence, we're following convention here.

Select line drawings of Scleromochlus taylori fossils from Benton (1999). There are two specimens here, showing dorsal and ventral views. The specimen on the right is the holotype, and the left shows two associated individuals. Note the banded scales crossing the vertebrae of the larger individual.

Although represented by at least seven specimens from the Lossiemouth Sandstone Formation, no Scleromochlus is well preserved (Benton 1999). Most specimens comprise shallow sediment molds rather than actual bones, and none are complete. But we should consider ourselves lucky we know of this animal at all: the delicate, 180 mm long bodies of Scleromochlus occur in sandstone deposits representing an ancient, wind-blown desert with 20 m high dunes. Such deposits are often devoid of fossil remains, but the Lossiemouth Sandstones actually preserve a diverse reptile fauna (Benton and Walker 1985). Still, it’s remarkable that the tiny bones of these reptiles preserved at all in these harsh conditions and in relatively coarse (fine - medium) sands - the grains preserving Scleromochlus are each as large as Scleromochlus teeth. As is typical of Lossiemouth Sandstone specimens, most Scleromochlus fossils are more-or-less articulated and many appear to have been crouching at death. With little indication of sun-cracking or scavenging, their remains clearly represent animals which were buried alive or buried shortly after death, probably by sandstorms or dune collapses (Benton and Walker 1985). Although likely complete when buried, no specimens have survived intact to the present. Cross-scaling elements from different specimens has permitted a reasonable insight into Scleromochlus anatomy all the same (Benton 1999). Some details remain murky however, and disagreement persists over precise bone lengths and skull bone attitudes (Sereno 1991, Benton 1999; Padian 2008). This is perhaps expected, given that Scleromochlus remains are interpreted via low-angle light and plaster or plastic peels of the skeleton molds. Bennett (1996) sums up working on Scleromochlus as "low-angle illumination [is used] to examine and interpret molds and peels, but in my experience a considerable amount of imagination is necessary as well".

Anatomy

Specific details aside, palaeontologists are happy to say that the basic bauplan of Scleromochlus resembles a small lizard with enormous hindlimbs (below). The skull has a low lateral profile but is rather triangular in dorsal aspect, with a blunt muzzle and widened posterior. So far as can be seen, the orbit is by far the largest opening in the skull, making the reduced nares look even smaller by comparison. The temporal fenestrae - as illustrated by Benton (1999) - are fairly sized, although their full margins aren't clear in any specimen. These sit above a posteriorly lengthened retroarticular process on an otherwise fairly unremarkable lower jaw. Each jaw seems to house 15/16 teeth, which are apparently isodont and - so far as can be seen - relatively small and lanceolate. The lizard-like visage of Scleromochlus is further enhanced by its short neck, which contrasts with later ornithodirans. The tail appears rather short too, being about as long as the snout-vent length.

Reconstructed skeleton of Scleromochlus taylori from Witton (2013), a modified version of the skeletal in Benton (1999).

The limbs of Scleromochlus are where a lizard-like visage starts to unstick. The forelimb bones are long and slender, and capped with tiny hands. The fingers are poorly known, but the tiny metacarpals suggest they were rather diminutive and unlikely of any use for standing or walking, a hypothesis supported by the dichotomy in fore- and hindlimb length. Even less lizard-like are the hindlimbs, which are extremely long - about half the length of the entire animal - and end with a narrow foot with tightly bound metatarsals. Both the forelimbs and pelvis appear relatively small compared to the legs, though neither is atypically small for the length of the animal. The fifth toe appears to have been lost, the only remnant being a short, pointed metatarsal. Scleromochlus hindlimb arthrology betrays a parasagittal posture akin to that of dinosaurs and pterosaurs - the suite of characteristics associated with this is one clue that Scleromochlus is closely related to these clades (Bennett 1996; Benton 1999; Hone and Benton 2008).

Thin, transversely-banded scutes(?) covered the dorsal surface of the Scleromochlus torso, extending from at least the shoulders to the posterior pelvic region (indicated in the fossil illustrations, above). These indicate that Scleromochlus was at least partly scaled, although whether this represents the entire integument is not clear. It is increasingly apparent that scraps of fossil skin do not tell whole stories about ornithodiran integuments, as more and more specimens with extensive skin preservation present 'mosaics' of scales, naked skin and various kinds of filaments (demonstrated in pterosaurs, theropods and ornithischians; e.g. Bakhurina and Unwin 1994; Chiappe and Göhlich 2010; Godefroit et al. 2014). Scleromochlus may have been covered in scales, but it is equally likely that it had fuzz-like filaments in places. There are several reasons for this. Firstly, it belongs within a phylogenetic bracket where filaments are the ancestral condition or, at very least, scales were prone to developing filamentous morphologies. Secondly, virtually all models of archosaur evolution recover Scleromochlus as sister taxon to a fuzzy clade - pterosaurs, so there is good 'phylogenetic proximity' for fuzz. Thirdly, insulating integuments are common - if not ubiquitous - in small, active (see below) desert-dwelling animals. Thus, while the overall  integument of Scleromochlus remains mysterious, a mosaic of filaments and scales is not an unreasonable suggestion. In the reconstruction here, Scleromochlus is shown as rather fuzzy all over (see below for rationale), with filaments poking through its scaly back as they do on opossum tails and armadillo hide.

Please provide your own 'boing' sound effects.

Locomotion

Lifestyle and palaeoecology

Spinosaurus, big and small versions, poking about a stream in Cretaceous Morocco. Someone's about to ask if these chaps should have humps or sails - head to Palaeontology Onlinefor my thoughts on this, and read on for why it doesn't resemble the (in?)famous National Geographic thumbnail.

Corking news, all: I've got a new article out at Palaeontology Online on the accuracy of palaeoart, explaining how confident - or not - we can be about different aspects of extinct animal appearance. It features two new bits of palaeoart, a Spinosaurus and a bizarre - but not implausible - reconstruction of Camarasaurus. The former started life as a simple illustration to make a point about reconstructing fatty tissues (including camel-like humps) in fossil animals, but I thought it warranted some elaboration: the painting above is the result.

The goal of the Palaeontology Online piece is not another 'how to?' guide to palaeoart, but a piece specifically targeted at those who want to know how accurate our restorations are. I've attempted to outline the reliability of standard palaeoart methods including phylogenetic bracketing, restoring musculoskeletal systems, placing fatty tissues, choosing integument types and, of course, deciding on colours and patterns. Note that the few years of optimism we've had for restoring fossil colour using melanosomes are over, because several new studies have highlighted numerous concerns with this technique: more on that at Palaeontology Online. All Yesterdays gets further mainstreamification, as does the mysterious, unexplained 'Support Original Palaeoart' logo (more on that in due time), and there's some philosophising over the goal of palaeoartists: are we actually bothered about 'the truth', or more concerned with making plausible art in line with fossil and biological evidence? OK, that's enough signposting for now: point your browser this way for the full piece, and be sure to leave any feedback below.

Just a quick note on the Spinosaurus illustrations here and in the article: they are not based on the thumbnail image of a unusual Spinosaurus skeleton at the National Geographic website, despite this spawning much excitement, umpteen new spinosaur renditions and revisions to Spinosaurus illustrations all over the Web. As stressed at Palaeontology Online, palaeoart is a scientific process requiring verified and trustworthy data. We have no idea how reliable the radical National Geographic depiction of Spinosaurus is because no information about the mount has been made public, and the image itself is tiny: it's silly to think there's enough resolution there to understand its anatomy. Moreover, there's enough counter-intuitive and weird morphology in that tiny photo to justify waiting for the data behind the mount to be published so it's accuracy can be evaluated. I'm not saying it's wrong, but I am joining the chorus of bona fide theropod experts in suggesting restraint against adopting it as the 'definitive new look' for Spinosaurus until we know more about it. The reconstructions here and at Palaeontology Online are based on Scott Hartman's skeletal: the appearance of the juveniles is speculative.

Coming soon (probably): exciting news from the world of sauropods!

I try to avoid hopping on the bandwagons following new discoveries - few internet experiences are more tiresome than seeing social media and inboxes swollen with discussions and pictures of the same new fossil species (tyrannosaurids, for some reason, do this more than anything else). Of course, some new discoveries are just too cool to pass up: Dreadnoughtus schrani Lacovara et al., 2014 is one of them. Not only does it have a fantastically marketable and charismatic name entirely befitting one of the largest land animals to ever exist (take that, naysayers), but the sheer amount of data published on it is really first class (Lacovara et al. 2014) and the fossil is truly spectacular. If you've not done so, check out the Dreadnoughtus description and supplementary material: there's everything from measurements and photographs to interactive 3D scans of every bone for you to look at in fine detail (or spin around like crazy while giggling, if you're comfortable enough with your maturity). And before you can say 'paywall', this is all freely-available, open access information. It's not just a great paper for those interested in sauropods or dinosaurs, but also an important reference point for those interested in the evolution of extreme animal anatomies and gigantism.

How the world met Dreadnoughtus schrani in palaeoart. Left, restoration by Jennifer Hall; right, Mark A. Klingler. Images from the Dreadnoughtus media release hosted at the Drexel News Blog.

I found one aspect of the very good, super-comprehensive and fittingly giant media release for Dreadnoughtus rather unusual, however: the artwork. For a media story principally being sold on the size of a dinosaur, the two 'official' pieces of Dreadnoughtus artwork by Mark A. Klingler and Jennifer Hall (above) have - what seem to me at least - some odd choices as goes composition and posture which might undermine the awesome size of Dreadnoughtus. I'm not saying the images are bad or 'wrong': there's lots of lovely detail and atmosphere in both (note the neat sauropod and titanosaur characteristics like the lack of manual claws, the concave posterior surface of the hand etc.), and this is not a dig at the artists, who have definitely earned the wide success of the Dreadnoughtus press campaign. My problem - and I hope this comes across as the constructive criticism it's intended as - is that I'm a bit underwhelmed by the sense of scale, which I'd say is pretty important for artwork of this animal. To be fair, conveying extinct animal size in art is never straightforward, but peculiar compositional choices in each image prohibit my being fooled into thinking I'm looking at truly giant animals. For example, both position the animals in the foreground, filling the canvas with as much Dreadnoughtus hide as possible. I can understand why - it says "it's so big we can barely contain it in the edges of these illustrations", but it also leaves little room for a point of size reference between us and the animals. It also forces the adoption of stooping postures and requires significant foreshortening to fit the animals into view, the former reducing their apparent size and the latter obscuring proportions we intuitively recognise as characteristic of large animals (e.g. the relatively small heads of large animals). Hall's illustration also sets the point of view at shoulder height so we're actually looking across and somewhat down at the subject animal - not necessarily what you might want to suggest this thing was bigger and taller than us. Both images feature trees immediately alongside their animals as a means of conveying scale, but I find the rest of the composition overpowers their effect. In all, while the other aspects of the images are effective, I'm just not sold on the size.

I find these decisions interesting because I think they represent a case of a modern palaeoart convention overruling 'classical' artistic approaches. Traditionally, artists use the same basic techniques for making subjects look big and important when placing them in a scene. They stress proportional extremes (including small head size - this even occurs in renditions of royal or divine human figures), use low points of view so that the the top of the subject clears the horizon line along with other elements in the composition, and place items to give an appropriate sense of scale. Positioning smaller items in the foreground can help the viewer find their position in the scene and ground their sense of size, but these need to be placed carefully: cluttered compositions tend to dwarf their subjects. A consequence of these methods is that giant subjects are often no closer than the mid-ground. An obvious exception to this are images with points of view positioned at the very base of a subject, looking up, so it looms above the viewer (below). This is a slightly different approach to the problem, though, almost treating the subject as the landscape rather than an entity within a background.

A cockroach-eye view of a titanosaur.

Palaeoart produced before the 1970s/1980s stuck to the classic rules of depicting giant animals: Zallinger, Knight, Burian et al. rarely deviated from 'standard' methods of conveying large size when drawing sauropods and other big extinct animals. The scientific transformation of dinosaurs into dynamic, active animals in the late 20th century also brought on a artistic shift where some artists abandoned 'classic' compositions in favour of more exciting, convention-defying and 'extreme' images. One consequence of this was some artists moving (frequently giant) animals closer to the foreground, turning them to face viewers and sometimes, through their body language, 'interacting' with those looking at them. The first seeds of this were probably sown by by the likes of Robert Bakker who, in many of his illustrations, fills every possible square inch with his animals to the point of using extreme postures - particularly arching backs and curving tails - to do so (e.g. illustrations in Bakker 1986). Bakker's works frequently lack the context of backgrounds however, leaving other artists to bring dynamically posed, big extinct animals closer and closer in landscaped works. I think Mark Hallett may have be particularly instrumental here, with works such as his famous 1984 'Dawn of a New Day', and the 1985 paintings 'Awakening of Hunger' and 'Ancient One' leaning towards, or perhaps even pioneering, an 'in your face' style of palaeoart where the subjects are looking at, sometimes menacing, their viewers (if anyone did this earlier, please let me know). Such artworks would become common in the 1990s, with Luis Rey famously combining these compositions with extremes of colour, perspective and pose to produce a style which has since been widely imitated. It's from such imagery that 'slasher' palaeoart arose, those images were animals are rushing, teeth and claws bared, at the viewer from within the painting.

Attitudes towards these foreground-emphasised, perspective heavy images are often divisive among palaeoart aficionados - some love them, others hate them. Fans of such works point out their utility for outreach, in that they're relatively novel, different, fun and striking, while detractors note their distortion of proportion, not to mention that many look, well, silly (I've argued elsewhere that this may have negatively skewed public perception of feathered dinosaurs). The most relevant common complaint to our discussion is that they lose all sense of scale, essentially for all the reasons listed above: unfamiliar proportions, a lack of foreground space to place 'scaling' elements, and often the loss of height associated with moving the anatomy into a position where it can all be seen behind the head (for many infamous examples, see Brusatte and Benton's enormous book Dinosaurs (2008)). Whatever your opinion, we can't deny their success and influence. such images are now a standard palaeoart convention, particularly in children's books, and have been used to showcase virtually any prehistoric animal you can think of. In this respect, the arching, frame-filling Dreadnoughtus images released last week are just following this now familiar palaeoart convention.

Thing is, I'm not sure if this practise works for all palaeoart, and especially in images where conveying size and anatomical details are important. Of course, the ultimate success of a composition is a matter of taste, and there is no actual 'right' or 'wrong' to palaeoart so long as it obeys basic laws of anatomy. But here's the beef: palaeoartworks often have a purpose - very commonly to convey the anatomy and size of a new species - but 'full frame' animal compositions are probably the worst composition to demonstrate these attributes, for reasons discussed above. Moreover, and fundamentally related to the goal of palaeoart being realistic portraiture of extinct species - how do we rationalise the adoption of the contorted postures required to fit the animals into frame? Why would these animals be condensing themselves into such weird shapes? And what do these poses look like from other angles? Wouldn't they look, at best, a bit odd? For me, seeing a restored animal in an unconventional, maybe even biomechanically implausible pose so it can take up more of the canvas is jarring, a reminder than I'm looking at an reconstructed animal rather than one an artist saw with their own eyes.


For art where proportions and a sense of scale is important, pushing our subjects back to the tried and tested middle distance would alleviate these problems, without jeopardising their excitement. Palaeoart was just as inspirational and exciting to audiences before we started rendering animals right under our viewer's noses, after all. Ultimately, while there's nothing inherently 'wrong' with any composition in palaeoart, some compositions suit certain scenes and animals more than others, and some are definitely more informative and educational than others. 'Full frame' compositions certainly have their place within palaeoart, but they're probably more limiting artistically and educationally than the alternatives.

I'll leave you with my own take on Dreadnoughtus, a quick painting done as the end result of my spate of fanboyism on Thursday night. And if you like sauropods, stay tuned, because there's more on the way...

The mighty Late Cretaceous titanosaur Dreadnoughtus schrani, making a mockery of two abelisaurids just by existing. Abelisaurids aren't known from the same formation as Dreadnoughtus, but are the most likely theropods to have occurred there given their abundance in the other Late Cretaceous South America. These are loosely based on Aucasaurus.

Update: 07/09/2014, well past bedtime

Not many moments after posting this, arty chum Jon Davies (@SovanJedi) responded with an image on Twitter which sums up the few thousand words above into one image:

The evolution of the dinosaur book cover: pic.twitter.com/ehszJQjS3J
— Jon Davies (@SovanJedi) September 6, 2014

It's funny because it's true.

References

• Bakker, R. T. (1986). The Dinosaur Heresies. London, Penguin.
• Brusatte, S. and Benton, M. J. (2008). Dinosaurs. Quercus.
• Lacovara, K. J., Lamanna, M. C, Ibiricu, L. M., Poole, J. C., Schroeter, E. R., Ullmann, P. V., Voegele, K. K., Boles, Z. M., Carter, A. M., Fowler, E. K., Egerton, V. M., Moyer, A. E., Coughenour, C. L., Schein, J. P., Harris, J. D., Martínez, R. D., and Novas, F. E. (2014). A gigantic, exceptionally complete titanosaurian sauropod dinosaur from Southern Patagonia, Argentina. Scientific Reports. 4, 6196; DOI:10.1038/srep06196.

Rukwatitan bisepultus a new titanosaurian sauropod from the Middle Cretaceous (Aptian/Cenomanian) of Tanzania. Why does this otherwise chirpy scene feature a dying Rukwatitan? Read on...

Hot on the heels of super titanosaur Dreadnoughtuscomes another new Gondwanan titanosaur, Rukwatitan bisepultus Gorscak et al. 2014 (press release restoration, above). As indicated by the publication of two new sauropods in close succession, we live in a time where our knowledge of sauropods dinosaurs is expanding rapidly. This surge in interest and activity is perhaps less conspicuous than other expanding areas of palaeontology - sauropods don't grab the headlines as much as small, feathered theropods - but it's fair to say that the landscape of sauropodomorph research has changed considerably in the last two decades. This particularly applies to our appreciation of their diversity and distribution across space and time. Once, the sauropod story could came to a near-end in the upper Jurassic once diplodocids started to decline, but we now know that titanosauriforms kept the sauropod end up throughout the Cretaceous, being abundant, widely distributed and diverse until the end of the Mesozoic. They appear particularly important in South America, where something like 39 species have been recovered (Gorscak et al. 2014).

Schematic of known elements of Rukwatitan bisepultus. From Gorscak et al. 2014.

Rukwatitan bisepultus is not South American however, but African, specifically from the 'middle' Cretaceous (Albain-Cenomanian) Galula Formation of Tanzania. Africa's Mesozoic faunas remain poorly known and, as one of only four named sauropods from 'middle' Cretaceous Africa, as well as a component of relatively poorly-known sub-Saharan Cretaceous forms, Rukwatitan is a find. Thus far, Rukwatitan is the only named sauropod from the Galula Formation, but other Galula fossils record a 'typical' Gondwana fauna of gondwanatherian mammals, notosuchian crocodyliforms (including the carnivoran-immitating Pakasuchus, below) and osteoglossomorph fish, as well as indeterminate small theropods and turtles (Roberts et al. 2010). Rukwatitan can be seen as another component of a middle Cretaceous sub-Saharan sauropod assemblage, joining the roughly contemporaneous sauropods, Malawisaurus dixeyi and Karongasaurus gittelmani of Malawi, along with scrappy fossils which hint at additional species. Represented by an incomplete skeleton (above) and a referred humerus, Rukwatitan is a relatively small titanosaur, although its exact size is difficult to gauge. It is seemingly larger than the relatively completely known (and probably closely related, see below) Malawisaurus, Rukwatitan humeri being 20 and 28% larger than those of Malawisaurus. With Malawisaurus estimated at about 9 m long (not 16 m as indicated in Paul 2010! - see comments below), this puts Rukwatitan in a rough length ballpark of 10-12 m.

Rukwatitan is not my first artistic trip to ancient Galula: in 2010 I helped Patrick O'Connor et al. restore their unusual notosuchian crocodyliform, Pakasuchus kapilimai, famous for it's cat-like slicing dentition. The word on the palaeo grapevine is that there's a lot more to come in the world of African crocodyliforms. I'd like to have another crack at rendering these guys, so I'll be waiting by the phone if anyone wants me...

Titanosauria is an increasingly big group, so leaving Rukwatitan with this label doesn't tell us much about its relationships to other sauropods. A useful phylogenetic landmark within Titanosauria is Lithostrotia, the group of derived titanosaurs which includes many famous taxa: Saltasaurus, Opisthocoelicaudia, Alamosaurus, Nemegtosaurus and Malawisaurus. This clade also contains all known armoured titanosaurs, although armour is not ubiquitous across the group (D'Emic et al. 2009). Other titanosaurs form successive offshoots from the titanosaur evolutionary line leading to Lithostrotia, and it's among these that Gorscak et al. (2014) place Rukwatitan. It only just misses inclusion within Lithostrotia however, suggesting close evolutionary ties to basal members of this group, including the geographically and stratigraphically proximal Malawisaurus. This mirrors findings that some geographically proximal, middle Cretaceous sub-Saharan reptiles - most notably Crocodyliformes - are also closely related, and substantiates ideas that sub-Saharan faunas were evolving at a relatively local, as opposed to cosmopolitan, or even continental-scale level (O'Conner et al. 2006; Gorscak et al. 2014). Possible further evidence of sub-Saharan regions being biogeographically distinct in the mid-Cretaceous stems from an apparent absence of many north African dinosaur groups. Although titanosaurs occur across the continent, evidence of large theropods (spinosaurids and carcharodontosaurids), other sauropod groups (rebacchisaurids, non-titanosaurian titanosauriforms) and ornithopods is currently lacking in Albian-Cenomanian deposits south of the Sahara. Will these animals turn up in time? Perhaps, but the continental Cretaceous beds of Tanzania and Malawi are not new localities only now being exploited, but the sites of many years, even decades of fieldwork. If north African dinosaur groups were there, their fossils are remaining well hidden.

Giving Rukwatitan a tighter address within Titanosauria helps us flesh out a rough projection of its bauplan with a little phylogenetic bracketing. The neck was probably relatively long compared to the tail, evidenced by phylogenetic neighbours and proportions of the preserved vertebrae (note that the schematic above is probably a little wimpy on the neck end of things). It's limbs were likely robust and relatively equally sized, and it's skin probably lacked osteoderms. A short, deep skull seems likely because Rukwatitan is bracketed by short-faced Titanosauriformes, but note that the bracket here is quite loose thanks to the deficit of sauropod skull material. We leant heavily on the well-known anatomy of Malawisaurus for our reconstruction (Gomani 2005), including Scott Hartman's skeletal.

Rukwatitan: river victim

Quarry map of the Rukwatitan holotype specimen, looking at the cross-section of the quarry stratigraphy rather than a 'birds eye view' of a specimen spread over a single horizon. Note the distribution of the skeleton over two layers, the mudstones (representing overbank deposits - the riverbank) and sandstones (fluvial deposits - the river channel). From Gorscak et al. 2014.

If you're moved by the story of Rukwatitan specimen RRBP 07409 and want to know how you can help dinosaurs who've suffered river attacks, please contact me for details of charities and fundraising events.

And finally, a request

Oh no, what caption to use? 'Pods of War? 'Podageddon? DinosAWESOME? Too... many... puns...

References

UPDATE: 21/09/14: Following chats with Nizar Ibrahim and Simone Maganuco, it appears the Spinosaurus 2014 saga has another twist to take concerning the controversy over the revised hindlimb proportions. I'd rather write about it in a comprehensive fashion when I have the time (hopefully tomorrow) and am hoping to deliver some definitive, knock-out information from the authors which puts this controversy to bed. Bear this in mind before you read the following...

ANOTHER UPDATE: 22/09/2014: Read this.

Are depictions of Spinosaurus like this now redundant? Answer: who knows? After weeks of anticipation and teased images, the 'new look'Spinosaurus has met a sceptical reception from academics and the online palaeontology community, and they've not kept their opinions quiet.

One thing is clear a week after the 'Spinosaurus reboot' (a phrase coined by Mickey Mortimer) was revealed amidst a furore of academic and media swirl: Spinosaurus c. 2014 has not met the warmest reception from the palaeontological community. A sceptical tone, sometimes very openly so, can be seen in numerous articles from the first popular science write-ups to articles penned by professional palaeontologists. As we all know by now, the primary concerns centre around Ibrahim et al.'s (2014) new Spinosaurus aegyptiacus reconstruction, which Brian Switek describes as a 'hodgepodge [of] different dinosaurs... the new subadult skeleton, digital representations of the original and long-lost Spinosaurus bones, vertebrae and hands that may or may not belong to Spinosaurus, as well as replacement parts from an assortment of spinosaurs'. Allegations have been made that scaling errors are responsible for the unusual new bauplan rather than an unprecedented lifestyle, with the allegedly tiny legs being far more proportionate once the scaling problem is addressed. These undermine the credibility of the furthest reaching claims of the authors - theropod quadrupedality and a lifestyle/locomotory strategy akin to early whales. Two widely shared and commented blogarticles on this topic over at Scott Hartman's Skeletaldrawing.com have cast enough doubt over the new reconstruction that the Spinosaurus 2014 authors publicly responded to the criticism, but the reply is really just a holding message. Other than pointing out well known problems of measuring images rather than fossils (which, to be honest, are unlikely to produce the large scaling problems levelled at the paper), the message is essentially 'all will be clear in an upcoming Spinosaurus monograph'*.

*For what it's worth, I took five minutes to measure up the new Spinosaurus skeletal restoration myself following Nizar Ibrahim's measuring instructions for dorsal vertebra 8, just to see if I could make head-or-tail of the debate. Differences in measuring landmarks were chalked up as being a potential problem, so I measured the ilium and femur blind to other methods, instead using whatever landmarks were most intuitive. For both the ilium and femur lengths, I arrived at almost identical scaling errors to Scott, and the legs should - according to the data in the paper - be c. 25-27% larger in the reconstruction. Something - the original measurements of the specimen or the reconstruction - just doesn't add up, and I suspect the latter, as I figure someone would have owned up to and corrected simple numerical errors in the paper by now. My working is below.

Independent test of the alleged hindlimb proportion issues in the new-look Spinosaurus. Skeletal reconstruction from Ibrahim et al. (2014); see Skeletaldrawing.com for the posts inspiring this test, especially this and this.

The controversy extends much further than just scaling, however. Acrossotherarticles, multiple issues have been raised including the incorporation of isolated spinosaur elements and other taxa to a single Spinosaurus reconstruction; whether all the material used in the reconstruction is of spinosaurid origin (e.g. this humerus); the likelihood for theropod quadrupedality (remember that we don't know anything concrete about Spinosaurus forelimbs: there is really nothing to suggest quadrupedality in this animal other than its alleged proportions); the authors taking too much credit for the 'semi-aquatic hypothesis'; the suitability of their journal choice and the somewhat ambiguous circumstances surrounding the provenance of the new material. And this is to say nothing of the extensive discussion on social media, much of which revolves around the same topics. This is not to say the Internet is hating on Ibrahim et al. (2014) - I think the pieces linked to here are balanced, reasoned critiques, not slanderous attacks - and, before anyone asks, I'm not saying I agree with, or even have opinions on a lot of these issues. The point here is that the 'Spinosaurus reboot' has experienced a very bumpy, almost slightly hostile landing.

The response to the Spinosaurus reboot is of some interest. Controversial, questionably-supported claims are made in palaeontology all the time, but they don't get the online palaeontology community anywhere near as riled as Spinosaurus has in the last seven days. Ibrahim et al. (2014) clearly hit a nerve, perhaps because they have inadvertently created a 'perfect storm' for scientific backlash.

At the heart of the storm is a data vacuum about Spinosaurus - an odd state to be in seeing as we're now meant to have a good idea what it looked like. The main discussion about Spinosaurus in the last week has been methodological: that is, trying to figure out how the new reconstruction has been put together. This is because the paper lacks essential details concerning how the 'hodgepodge' of spinosaur bits were scaled to size or identified as Spinosaurus aegyptiacus in the first place. In skipping these details readers are left guessing - and discussing - how the proportions were ascertained and whether they are trustworthy. That people would want to know this was predictable: you can't propose a radical notion like a famous theropod being a semi-aquatic quadruped, even converging on whale ancestors, without academics, enthusiasts and dinosaur nerds wanting to know more. While the paper does have plenty of good data, it lacks transparent methods and discussions where it counts, leading readers to make their own tests and discoveries. Lest we forget, people like talking about dinosaurs online at technical levels, and it's only natural that blogging software and social media is being fired up to discuss these revelations. It's quite likely that there'd be less fuss made if the paper stood on sounder methodological ground but, ultimately, controversy sells, in part because the continual uncovering of new information and scientific debate makes for good copy.

Compounding this effect is the star of the show: Spinosaurus itself. By now, Spinosaurus has to be one of the most popular dinosaurs of all. It's the one widely known theropod to have a size advantage over Tyrannosaurus, has starred in a couple of big movies and documentaries, is undeniably cool looking, is a bit 'alternative' as dinosaurs go... for lots of reasons, it's a major dinosaur celebrity. Even among po-faced academics, the sheer size and unusual anatomy of Spinosaurs means most - probably even guys who work on brachiopods - find it a little bit more interesting than usual. Any publication on this animal is guaranteed a good amount of casual interest, but one where the animal is almost completely reinvented will send the online palaeontology community into overdrive. Did anyone else have to wait for the Science website to stop crashing when the embargo was lifted last week? I'd be interested to see how riled the internet palaeontology community got if someone questionably reconstructed a small ornithischian. For contrast, consider that the publication of another dinosaur with a radical lifestyle - the burrowing dinosaur Oryctodromeus - ruffled relatively few feathers when it was published, despite it's PR. I remember most discussion of it on the Dinosaur Mailing List concerning the formulation of its name.

Driving the storm is the considerable hype surrounding the paper, which bears little resemblance to traditional scientific press releases and is more akin to the launch of a summer blockbuster. 'Surrounding' is the right word, too, as tantalising glimpses of the new reconstruction were online weeks before the paper's release, foreshadowing the avalanche of 'official' art, articles, and videos which would follow. There are documentaries, a tie-in exhibition in Washington DC, press conferences and lectures. You'd think Spinosaurus and its wranglers were rock stars. I mean, can you name one other palaeontological PR event which needs dry ice?

The popular side of this release has been a resounding success, which - whatever you think of science being spun as a media event of this kind - is certainly well earned. In concert with National Geographic, Ibrahim et al. (2014) have put on a very slick, professional show with some wonderful art and graphics, and they've certainly made it difficult to miss. But publicity can be polarising, not to mention difficult to steer. It seems the PR for Spinosaurus 2014 has somewhat backfired in the palaeoblogosphere, the conspicuous, sensational nature of the story encouraging interested minds to investigate and test, and ultimately question the findings at the core of the hype. I expect the extensive publicity surrounding a widely-questioned paper also brings a faint sense of irritation to some, prompting them to advertise the fact that the conclusions are not as watertight as the documentaries, exhibition and magazine covers indicate. Whereas other studies with problematic conclusions would slip away into the literature to be discussed within the closed confines of scientific journals, Spinosaurus 2014 cannot hide easily: the advertising and publicity for this paper is keeping the controversy relevant and prompting more responses. I do wonder what National Geographic, presumably footing the bill for all this press work, are making of the frosty scientific response to Spinosaurus 2014.

Between the data vacuum of a radical new proposal, a megastar fossil animal and persistent reminders of a controversial study, it's hardly surprising that the online palaeo community has spent the week giving the Spinosaurus reboot a good grilling. What does the future hold? With the promise of a Spinosaurus monograph, we can be sure that there will be more discussion eventually, but, more realistically, the next major ripples will follow response papers. Some authors are already in talks about this and - given what's been demonstrated online already - there are strong cases to be made against the main hypothesis of the Spinosaurus reboot. Is a rebuttal article appropriate with another paper on the way? Yes, entirely, because we have to work with data which is available and test the hypotheses presented to us. In this case, the new-look Spinosaurus and the many implications made about its habits have been quickly questioned - deemed irreproducible, even - by a number of scientists, and this should be 'formalised' as a genuine concern about the initial paper. The upshot, of course, is that the eventual monograph will have to take this into account, which should make for a stronger publication, and hopefully an improved understanding of Spinosaurus itself.

I can't help think that there are a few causalities from the last week, not least being the good new data in Ibrahim et al. (2014), such as Spinosaurus weirdly tetradactyl feet, unusually short femur and dense bones (Ibrahim et al. 2014). What do these mean, in light of the hindlimb scaling controversy? Is the long first toe more to do with spreading weight than creating a flipper? Are the thickened bone walls more to do with relocating the centre of gravity than swimming? There are interesting discussions to be had there, but they've been overshadowed by other details. Also, scaling issues or not, I imagine the 'dachshund'Spinosaurus is here to stay for a while, so we can look forward to having to downplay confidence about the new reconstruction of Spinosaurus for the foreseeable future. It's very doubtful that the press will be interested in a story about the uncertainty over a new paper, nor is National Geographic likely to replace the legs on its Spinosaurus model with question marks. This is a constant bugbear of working within science of course: the media is interested in new and exciting discoveries, but has virtually zero attention span for scientific debate.

Finally, is there anything to learn from this? For me, the message is that while publicity is largely about presenting conclusions and results, we can't just assume our audiences are passive. Particularly if you're discussing a fan-favourite species (and let's face it, 'fans' here includes a good number of vertebrate palaeontologists), people remain just as interested in what you've done as what you conclude, and omitting those details leaves papers, and those associated with them, vulnerable to misunderstandings and criticism. As demonstrated this week, even the combined might of Spinosaurus and its PR campaign is not immune to this: when the world's largest theropod took a bite out of the Internet, it was bitten right back.

Reference

• Ibrahim, N., Sereno, P. C., Dal Sasso, C., Maganuco, S., Fabbri, M., Martill, D. M., Zouhri, S. Myhrvold, N. & Iurino, D. A. (2014). Semiaquatic adaptations in a giant predatory dinosaur. Science, 1258750.

No-one with an interest in Mesozoic reptiles will have missed the week of controversy following Ibrahim et al.'s (2014) new reconstruction of Spinosaurus. The most important debate has focused on the allegedly reduced Spinosaurus hindlimbs, which are integral to the proposed locomotor and lifestyle hypotheses proposed for the 'new look' animal, but also difficult to reconcile with presented data. Scott Hartman, who's no stranger to producing high-quality skeletal reconstructions, blew this whistle first when he found the reconstructed proportions of the Spinosaurus neotype specimen - a series of vertebrae and hindlimb elements - were questionably scaled against measurements of the bones themselves. Lead author of the Spinosaurus study, Nizar Ibrahim, publicly responded and suggested that the measuring landmarks Scott used in comparing vertebral and hindlimb elements may be wrong. When reviewing the controversy before the weekend, I attempted my own scaling effort, using Nizar's suggested landmarks, but ended up replicating Scott's results almost exactly. I concluded "[s]omething - the original measurements of the specimen or the reconstruction - just doesn't add up, and I suspect the latter, as I figure someone would have owned up to and corrected simple numerical errors in the paper by now."

It turns out that I've got to eat a few of those words. Following my post, Nizar opened a chain of correspondence where I directly asked about these scaling issues. Nizar's response was bringing his coauthor Simone Maganuco into our chat, who had taken the time to demonstrate and describe how the restored vertebral and hindlimb lengths match the dimensions reported in the paper. In his screenshot and email, Simone provided an enlarged view of the restored Spinosaurus trunk and took the time to explain where he thought the alleged scaling errors came from. Appreciating their interest to a wide audience, Simone has kindly allowed me to reproduce his screengrab and email here.

Image courtesy Nizar Ibrahim and Simone Maganuco, used with permission.

Dear Mark,

It is nice to be in touch with you. I am writing to comment briefly on my photoshop image, forwarded by Nizar a couple of hours ago.

I hope it is the key to understand the misunderstanding about the measurements, so I would be really glad to know your opinion about it.

I have tried to replicate the coefficients for scaling obtained by you and Scott Hartman and here is my line of reasoning.

Look at the vertebra D8 in my photoshop image. For convenience, we can focus our attention on the D8 on the left.

The yellow line is 18 "units" (and matches our measurements in the table) but if you include the posteriormost margin of the slanted posterior face and the condyle you have nearly 23 units.

23:18=X:71 where 18 and 71 are also the measurements in cm in the table of the Science paper; 23 units is the length of the whole vertebra in the drawing; and X should be the length of the ilium to match the length of the vertebra in the drawing, if one assumes that the whole vertebra - and not the yellow line - is 18 units, i.e., if one thinks  we used different landmarks and measured the maximum length of the centrum.

The value of X is 90.72  units.

90.72 /71  = 1.27 that is exactly the coefficient for pelvic girdle and hindlimb scaling suggested by Scott @ skeletaldrawing.com to resize the pelvis and the legs to match the size of the D8 vertebra measured with different landmarks (i.e., if 18 is considered the maximum length).

I can see that your coefficient is slightly lower, and I wonder if you have taken slightly lower measurements (it seems to be the case looking at the white lines in your test).

Do you think that this could be the explanation of  what happened?

In the paper, we thought it was better to measure the vertebrae from rim to rim (the rounded margins of the faces), excluding the condyle, and at the same dorsoventral height (because some vertebrae are like parallelograms). It is easier to compare anterior dorsals and posterior dorsals in this way, and it is easier also to compare the centra with those of some specimens not prepared three-dimensionally but preserving well-articulated vertebrae, i.e. specimens in which it is difficult to look at the anterior condyle.

As what concerns the femur, it must be taken into account that there is also a slight perspective effect, because in the digital model it points a bit laterally. i.e., it is not 100% parallel to the sagittal plane.

The misunderstandings generated by the comparison between the figure and the table clearly indicate that we had to indicate our landmarks in one extra figure, or dedicate a couple of lines to this into the text to satisfy the need to compare figure and measurements by people who want to test our skeletal reconstruction.

When I work with palaeoartists to prepare illustrations and flesh-models I also compare figures and measurements, so I can understand this need.

Sometimes there are figures that are not 100% in the view indicated in the caption (also because it is not easy to put a bone in plane!) and sometimes it is difficult to understand the landmarks used to take measurements. What if I were in your shoes? Who knows... but I can understand that the new look of Spinosaurus has unexpected proportions that leads to think that there is something wrong.

In the monograph everything will be more clear because the detailed figures will report measurements directly on the bones, permitting everybody to see the landmarks.

In the meantime, however, I think it is useful to clarify this aspect.

Best wishes,

Simone

--

So there we have it: the measurements, landmarks and an image where they can be measured accurately. The latter is especially important because dorsal vertebra 8 in the full restoration is rather small, and thus prone to measuring errors even when measuring landmarks are known. A slip of a few pixels may not seem like much but, because the bone is a tiny component of a huge reconstruction, such minor errors can throw a scaling calibration right off. These risks were identified in Scott's originalposts, and it seems they have been borne out. Nevertheless, it is interesting that Scott and I - and others, according to some Facebook chat - found such similar results: this could be coincidence, or it might be that the published reconstruction lends itself to a erroneous interpretation. Either way, there is plenty of food for thought here as goes presentation and reading of reconstruction data. For the record, when attempting to replicate the scaling again, this time on the screenshot, I found my results matched measured values given in Ibrahim et al. (2014) within a few percent. My confidence in the published proportions is thus fully restored.

Hopefully this helps resolve the scaling controversy with the 'Spinosaurus reboot', and the result is much more confidence about the downright weird and remarkable anatomy of this genuinely unusual animal. Thanks to Nizar and Simone for taking the time to explain their work, and allowing me to post their response here.

Reference

• Ibrahim, N., Sereno, P. C., Dal Sasso, C., Maganuco, S., Fabbri, M., Martill, D. M., Zouhri, S. Myhrvold, N. & Iurino, D. A. (2014). Semiaquatic adaptations in a giant predatory dinosaur. Science, 1258750.

Quick sketch of Deinonychus antirrhopus with expanded, bone-puncturing jaw muscles, a requirement of having a bite as strong as a modern alligator. Say what? Read on...

There's a part in Michael Crichton's Jurassic Park novel where Velociraptor attempt to bite through bars to reach a people-shaped lunch. Presumably, they're meant to give readers something to rally behind seeing as one person in the line of fire is Ian Sodding Malcolm - I'd be chewing through steel too if it meant we could enjoy a few moments without another preachy monologue. Crichton describes them as hyena-like in their ability to bite through steel, delivering thousands of pounds of pressure per square inch and gnawing their way through thick metal bars in 15 minutes.

Dromaeosaurids biting through steel bars - heck, any animal biting through steel bars, including hyenas - intuitively sounds like crazy talk*. But was Crichton at least right about the strong bites of dromaeosaurids? I've been doing some investigating on dromaeosaur jaw muscles for a new palaeoart commission, but I've come unstuck. Here's why.

*Is there any substance to claims about modern animals biting through steel? Given that tooth enamel is only very slightly harder than straight steel, I wonder how long teeth would last when gnawing through anything but the thinnest metal sheet.
Recently, Gignac et al. (2010) presented a suite of bite marks on Tenontosaurus bones argued to show Deinonychus as capable of deeply puncturing bone with powerful bites. The tooth gouges match those of large Deinonychus in many aspects (bite mark size, shape, correspondence with dental arcade) and broken teeth associated with the same Tenontosaurus corroborate suggestions that Deinonychus fed from the carcass. Other teeth, not from Deinonychus, were also at the site, but their owner does not seem to have left any other obvious traces. Experiments with modern cow bones suggest Deinonychus needed a whopping 8200 N to puncture Tenontosaurus bones to the degree seen in the fossil remains. This value puts Deinonychus bites on par with those of adult alligators and leaves hyenas in the biomechanical dust. It also grants Deinonychus one of the highest estimated bite forces of any dinosaur, even greater than animals of much larger size. The tooth marks only match the largest known Deinonychus individuals, possibly indicating that juveniles were incapable of delivering such bite forces. Because Deinonychus puncture wounds are rare, Gignac et al. argue that puncturing bones was not common in Deinonychus, and that their powerful bites were primarily used for aggressive behaviours instead.

Bitemarks in the radius of Tenontosaurus specimen FMNH PR 2261, below, compared to the dental arcade of Deinonychus antirrhopus, above. This is one of many pathologies on FMNH PR 2261, almost all of which have been attributed to Deinonychus feeding behaviour. From Gignac et al. 2010.

For artists, Gignac et al.'s paper has important implications. Generating 8000 N of bite force requires a lot of muscle, so we might predict that Deinonychus jaws had the same swollen jaw muscles of modern crocodiles to generate all those bone-smashing newtons. This is at odds with other reconstructions of Deinonychus, where the jaw muscles do not atypically alter the contours of the face. I don't know how visible expanded, crocodile-like jaw muscles would be on deeply feathered maniraptorans, but reconstructions with sparse or naked faces would certainly need to take this on board. I've had a quick play about with this concept in the conservatively feathered Deinonychus above.

Problem is, Gignac et al.'s conclusions are not uncontested. Biomechanical assessments of Deinonychus jaws have found they were mechanically weak and ill-suited to delivering powerful bites (Therrien et al. 2005; Sakamoto 2010; Fowler et al. 2011). Therrien et al. (2005) estimated Deinonychus bite force at a relatively wimpy 15.7% of alligator jaw power, which Gignac et al. translate into 1450 N. This isn't unimpressive - as strong as that of a 30 kg wolf - but a far cry from an alligator-like bite, and certainly deflates our reconstructed jaw muscles to their traditional size. On the face of it, I certainly find the arguments for weak jaws more convincing. Hyenas and alligators have robust, wide and solidly-built skulls with generous room for jaw muscle placement, whereas the skull of Deinonychus is full of holes, is relatively narrow and slender, and with comparatively little room spaces jaw for muscles.

So, what to do? Jaws with relatively small muscles have been the norm in Deinonychus palaeoart since its discovery, but is it time we changed that? Were their jaws actually visibly and powerfully muscled as inferred by their trace feeding evidence, or is there something missing here? Is it significant that lower estimates of their bite forces match those of animals which can also puncture bone (wolves - see Haynes 1982)? If anyone has anything to add, please let me know...

References

The industry of reconstructing extinct animals in illustration, sculpture and animation - we all know it as 'palaeoart' - is a paradoxical place. One the one hand, there is more demand for palaeoart than there ever has been, increasing recognition of the role of palaeoart as a scientific and outreach tool and, because of the internet, more interesting and thought-provoking palaeoart being produced than ever before. This would make it seem that palaeoartistry is a flourishing, economically viable and interesting place to work within. On the other hand, much of our widely published, well-paid and/or high profile palaeoart work is rife with plagiarism, is creatively stagnant, has limited commercial appeal and presents gross inaccuracies to the fossils it is meant to represent. Given the elevated public influence and larger economy of these high profile artworks, it might be argued that this less interesting, ethically-questionable and scientifically dubious side of palaeoart overrides the independent sphere as the current 'status quo' within the palaeoart industry.

You guys are hypocrites. You've asked for/given free art, for instance, and been involved with products featuring awful palaeoart

So what can we do?

For various - and mostly good - reasons, there's not been much chance for blogging of late, but the upside is that I have a lot of new art, discussion and science to share in the near future. In the interests of not completely abandoning the blog in the interim, here's a series of Tweets posted over the last two days documenting work on a painting of the Early Jurassic pterosaur Dorygnathus banthensis. I can't say too much about the painting at this stage, because it's earmarked for an upcoming project and its context will be best explained there. Still, there's no harm in leaving a few notes about the restoration and painting process, so here goes...

Just for fun, I'll be posting progress on a painting of the early pterosaur Dorygnathus as I work. Here's the start. pic.twitter.com/5SWIkB1NzX
— Mark Witton (@MarkWitton) October 17, 2014

The initial digital sketch, complete with Kevin Padian's (2008) Dorygnathus skeletal reconstruction in the top corner for basic guidance. Padian's (2008) work was my principle reference here, and is probably the go-to paper for all things Dorygnathus. Those of you who know a bit about pterosaur research may be aware that Kevin was the main, and rather vocal, proponent of pterosaur bipedality in the 1980s and 1990s, so may be surprised to see his name attached to a quadrupedal pterosaur skeletal. Kevin is now on board with the consensus view that pterosaurs were primarily or exclusively quadrupedal animals, although he still argues that bipedality was essential for rapid terrestrial locomotion. I don't really agree with him, but that discussion will have to wait for another time.

90 minutes in, and Dorygnathus is starting to take shape. But it's tea time, so he'll have to wait for more work. pic.twitter.com/xSutwXrvYw
— Mark Witton (@MarkWitton) October 17, 2014

A little rotation of the underlying sketch, some basic outlines of the background complete the overall composition. I've had this image knocking about my brain for about a week now, and think the layout is a fairly good approximation of what I've been imagining. This painting has a message to deliver about the sprawling posture of the animal, and I think this composition demonstrates that well enough. There is a lot of compelling anatomical evidence that Dorygnathus and many other non-pterodactyloids could not adopt erect forelimb postures, which is partly why they're considered inferior terrestrial animals to pterodactyloids. But is that the case for all non-pterodactyloids? I'm saying nothing else at this stage, other than that this painting has a contrasting sister image.

End of play on Dorygnathus today. Tomorrow's goal: make him look 50% less evil. pic.twitter.com/lSPmgsURHp
— Mark Witton (@MarkWitton) October 17, 2014

A lot more detail by the end of day one. The eye was shrunk to fit the orbit a little better, and the animal now looks generally larger as a result. This is good: Doryngathus is about 1.8 m across the wings, so needs to look seagull-sized. The basics of the colour scheme are added now too. There's a lot of evidence that rhamphorhynchines* like Dorygnathus were seabird like in their habits, so it makes sense to use common seabird colours - whites, greys, blacks and - here. There's a butt-tonne more detail here than I'm used to working with, the result of a big upgrade to my painting hardware and software. A graphics tablet built this decade? Imagine that!

*I don't really agree with Bennett's (2014) proposal that Dorygnathus is a scaphognathinid/ine/whatever. Ah, non-pterodactyloid pterosaur taxonomy: what a mess.

Back on the Dorygnathus wagon. Starting work on the background: weathered limestone coastlines for the win! pic.twitter.com/fY4q9Pzwf7
— Mark Witton (@MarkWitton) October 18, 2014

Lots of laminae - fine scaling bedding - in the rock here. Got to put that training into sedimentology to use somewhere.

Quick Dorygnathus update before lunch. Not much more to do now... pic.twitter.com/ShltDx5krS
— Mark Witton (@MarkWitton) October 18, 2014

Nearly there by this stage. Note the similar dip-direction on the rocks jutting out into the sea. Their angle means we can have a few splashy waves here and there, which is nice, and you could map the geology of this bay quite effectively. Because if you had a time machine and visited the Jurassic, mapping grey rocks would totally be the thing to do.

And I think we're done. Dorygnathus on a rock, next to the sea, surrounded by the guano of his friends. pic.twitter.com/47pKhyXYYP
— Mark Witton (@MarkWitton) October 19, 2014

And done. The only real differences between the last two images are some tidier shading, a few background Dorygnathus and some splats of guano on the hero rock. I've long thought that locations supporting lots of pterosaurs would literally be a bit crappy, but never put it into art until now. I expect their guano looked a lot like that of birds and other reptiles: a mix of white, pasty stuff and darker gunge. Nice.

OK, time at the blog. Sorry for the short post, but I may have some good news soon for anyone interested in buying prints of my stuff - just in time for Christmas! I'll leave you with a larger version of the image than the low-res versions afforded by Twitter.

Dorygnathus banthensis at the coast, surrounded by the filth of its contemporaries.

Reference